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ISSN No. IXW3-7903

New Zealand Oceanographic Institute Memoir 106

Athecate Hydroids and their Medusae
(Cnidaria: Hydrozoa)

Peter Schuchert

The Marine Fauna of New Zealand:

ISSN 0083-7903,106 (Print)
ISSN 2538-1016; 106 (Online)

COVER PHOTO: Staurocladia wellingtoni, a new crawling medusa from Wellington'
south coast, diameter approximately 5 mm.

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NATIONAL INSTITUTE OF
WATER AND ATMOSPHERIC RESEARCH LTD

The Marine Fauna of New Zealand:
Athecate Hydroids and their Medusae
(Cnidaria: Hydrozoa)

PETER SCHUCHERT

New Zealand Oceanographic Institute
National Institute of Water and Atmospheric Research (NIWA)
P.O. Box 14-901, Kilbirnie, Wellington
NEW ZEALAND

Present address

Zoological Institute, University of Basel,
Rheinsprung 9, CH-4051 Basel
SWITZERLAND

New Zealand Oceanographic Institute Memoir 106

1996

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Cataloguing in publication:

SCHUCHERT, PETER

The marine fauna of New Zealand: Athecate hydroids and their medusae (Cnidaria : Hydrozoa)
Peter Schuchert — Wellington : New Zealand Oceanographic Institute, 1996
(New Zealand Oceanographic Institute memoir, ISSN 0083-7903, 106)

ISBN 0-478-08377-7

L Title: II. Series

Series Editor: Dennis P. Gordon

Typeset by: Rose-Marie C. Thompson

National Institute of Water and Atmospheric Research Ltd

Printed and bound for NIWA by
Colorgraphic International
A Division of Wilson & Horton Print Ltd
Printers

Wellington & Auckland

Received for publication
31 May 1995

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CONTENTS

Page

ABSTRACT 5

INTRODUCTION 6

METHODS ......ri..*.:%+■* 9

STATION LIST

SYSTEMATICS

Subclass Anthoathecata
Order Filifera

Family Clavidae .

Family Calycopsidae
Family Bougainvilliidae ..
Family Hydractiniidae
Family Rathkeidae
Family Cytaeididae ....

Family Pandeidae .

Family Protiaridae .

Family Eudendriidae
Family Proboscidactylidae
Family Eucodoniidae nov.

Order Capitata

Family Polyorchidae . 90

Family Zancleidae . 92

Family Cladocorynidae . 96

Family Porpitidae . 98

Family Corymorphidae . 101

Family Tubulariidae . 106

Family Boeromedusidae . 115

Family Margelopsidae . 116

Family Candelabridae . 117

Family Corynidae . 118

Family Cladonematidae . 131

Family Solanderiidae . 139

Family Pennariidae . 142

ACKNOWLEDGMENTS . -----.... - -. 144

APPENDIX

144

REFERENCES

145

INDEX

157

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Frontispiece:

Solmideria ericopsis (Carter), height approximately 55 cm, from 5 m, Riko Riko Cave, Poor Knights Islands.

Photographed by Dr C.N. Battershill.

The Marine Fauna of New Zealand:
Athecate Hydroids and their Medusae
(Cnidaria: Hydrozoa)

PETER SCHUCHERT *

New Zealand Oceanographic Institute
National Institute of Water and Atmospheric Research (NIWA)
P.O. Box 14-901, Kilbirnie, Wellington
NEW ZEALAND

ABSTRACT

The 79 species of athecate hydroids and Anthomedusae known from New Zealand are surveyed. Their
systematic revision is based on preserved museum material, newly collected living material, and life-cycle
observations. Three new genera, seventeen new species, and fourteen new records are described. Full definitions
are given for all taxa and almost all species are illustrated. One new family, Eucodoniidae, is defined and
incorporated into the order Filifera. The new family includes only Eucodoniuw brownei Hartlaub, 1907. The
three new genera are Fiordlnndia (Hydractiniidae), Barnettia (Pandeidae), and Fnbiennn (Proboscidactylidae).
The genus definition of Zmiclen is widened to include Zmiclella and Hnloconjne. The new species are: Barnettia
caprai, Boiigainvillia dimorpha, Corymorpha intermedia, Coryne tricycla, Ectopleura multicirrata, Eiidendriiun
maoriamts, Euphysa problematica, Eabienna sphaerica, Fiordlandia protecta, Hydractinia novaezelandiae, H, rubricata,
Merga treubeli, Podocoryna australis, Rhizogeton conicum, Staurocladia wellingtoni, Stylactaria otagoensis, and Z anclea
polymorpha. Two possible new species of the family Corynidae are not named specifically due to insufficient
life-cycle information. The life cycles of Bougaiiwilla vervoorti Bouillon, 1995, Boiigainvillia dimorpha n. sp.,
Podocoryna australis n. sp. and Zanclea polymorpha n. sp. are described. Ascidioclava parasitica Kirk, 1915 is
referred to Bythotiara parasitica n. comb, and its life cycle partially revealed. Cladonema novaezelandiae Ralph,
1953 is referred to Cladonema radiatum Dujardin, 1843. Titbiclava rubra Farquhar, 1895, Corydendrium rubra
Stechow, 1924, and C. zelandicum Stechow, 1924 are recognised as new synonyms of Turritopsis nutricula
McCrady, 1857. Coryne tenella Farquhar, 1895 is redescribed from its type locality and synonymised with
Sarsia eximia . Based on the examination of new living material of Solanderia ericopsis (Carter, 1873), the genus
Chitina Carter, 1873 is again sunk into synonymy with Solanderia Duchassaing & Michelin, 1846. Pochella
oligonema Kramp, 1955 is referred to Eabienna oligonema n. comb. Life-cycle observations were made on other
species to confirm their identity. New records are: Calycopsis bigelowi Vanhoeffen, 1911, Dicoryne conybearei
(Allman, 1864), Amphinema dinema (Peron & Lesueur, 1809), Halitholus pauper Flartlaub, 1914, Cladocoryne
floccosa Rotch, 1871, Ectopleura crocea (L. Agassiz, 1862), Coryne pusilla Gaertner, 1774, and Sarsia japonica (Nagao,
1962). Several other possible new records were not assigned to a species due to insufficient lifecycle information.

Keywords: Anthomedusae, Athecata, systematics, life cycle, review, revision, new species, New Zealand,
marine fauna

* Present address:

Zoological Institute, University of Basel, Rheinsprung 9, CH-4051 Basel, Switzerland.

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INTRODUCTION

General Remarks

The detailed studies of Ralph (1957-1961) made the
thecate hydroids of New Zealand comparatively well
known. In contrast, the hydromedusae and the athe-
cate hydroids are much less known. Kramp (1965,
1968) and more recently Barnett (1985) and Bouillon
(1995) have made major contributions to our know¬
ledge of the hydromedusae of New Zealand, whereas
investigations on most groups of athecate hydroids
are still fragmentary. Short reviews were provided by
Farquhar (1896), Bale (1924), and Ralph (1953). Ralph's
review also dealt with the history of earlier investi¬
gations on New Zealand Athecata and their medusae.
Since Ralph's (1953) study only few published con¬
tributions have been made. Hand (1961) described
Podocoryna bella, more information was published on
the peculiar hydroid Pelagohydra mirabilis (Pilgrim
1967a, 1967b; Rees & Ralph 1970), Watson (1987)
reviewed the Eudendriidae, Bouillon and Cornelius
(1988) redescribed the endemic Chitina ericopsis, and
recently Cairns (1991) revised the Stylasteridae and
described many new species.

A complete review and revision of the athecate
hydroids and their medusae was therefore much
needed and the present work aims to provide this at
least in part. It was intended from the very beginning
that this study should be written not only for the use
of specialists, but also offer other biologists, e.g.,
ecologists, specialists of other groups, and health
officers, a tool for identifying the known New Zealand
species. Therefore, almost all the species are figured
and described, whenever possible from living samples.
Where known, information on the life cycle is given
and occasionally figures from the literature have been
reproduced, especially where these publications are
difficult to obtain for the average institution. This was
done to allow identifications of life-history stages so
far not known from New Zealand. An identification
key for all the species is not provided. The reasons for
this are: many life-history stages are still unknown,
for many species complete life-cycle information is
necessary, and more species than recorded here can
certainly be found in New Zealand. A general key
would therefore in many cases be not applicable or
lead to an incorrect identification. As almost all species
are figured here, the beginner will be able to go quickly
through all illustrations and find a suitable taxon for
an animal to be identified. Diagnostic characters of

the New Zealand taxa are provided below the
family level. They are intended to highlight important
diagnostic characters.

The taxonomic range treated in this work does not
include one complete monophyletic group. This is due
to the exclusion of the Stylasteridae, which have
been revised recently (Cairns 1991), and the restriction
to marine or brackish-water animals. The Probosci-
dactylidae, which some authors include in the Limno-
medusae, are here included in the Anthoathecata
(Anthomedusae and Athecatae). Other marine Limno-
medusae are not known from New Zealand (for the
freshwater genus Craspedacusta, see Fish 1971). With
the exception of the above-mentioned Proboscidac-
tylidae, the overall taxonomic system adopted in the
present work mostly follows the system of Bouillon
(1985a) for the Filifera and Petersen (1990) for the
Capitata.

During the last decades, many efforts have been
made by hydroid taxonomists to overcome the sepa¬
rate systematic treatment of the polyp and medusa
stages and to present a unified system (e.g., Naumov
1969; Brinckmann-Voss 1970; Bouillon 1985a; Peter¬
sen 1990). Although many systematists, mostly for
practical reasons, still continue to work with one
stage only, such a separation of two life stages is very
unbiological. Additionally, especially in the Antho¬
athecata, only information on the complete life cycle
is often sufficient to identify species (cf. Sarsia and
Dipurena species). In accordance with other authors
(e.g., Edwards 1973b; Boero et al . 1992), the polyp is
seen here as a second larval stage (for an opposing
view see Cornelius 1990). There is nothing particular
in its ability to propagate vegetatively as this is also
well known in other invertebrates, e.g., parasitic
Platyhelminthes. Therefore, the life cycle of the hydro¬
medusae is not exactly a metagenetic cycle with
alternating vegetative and sexual propagation of an
adult (cf. discussion in Cornelius 1990). In this work,
both medusa and polyp stage, and also young medusa
stages, are presented together to underline their
biological unity, and many efforts were made to
elucidate life cycles.

The work of Petersen (1990) has clearly shown that
many hydrozoan taxa are para- or even polyphyletic.
This is certainly not only true for the Capitata exam¬
ined by him. A completely new system, based on
phylogeny, is very much needed and the revisions
made by Petersen (1990) are a desirable step towards

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that goal, but not all of his proposals can be followed.
One problem in hydrozoan systematics is the relative
simplicity of the animals combined with their pheno¬
typic variability. A cladistic analysis will always rely
on few and simple characters and therefore present
very tentative phylogenies. An analysis of phylogenetic
relationships using sequence comparisons might help
to resolve many ambiguities.

Many hydrozoans, and especially most athecate
hydroids, are rather inconspicuous animals. It is
therefore not surprising that their ecology is rather
poorly known (cf. Boero 1984; Gili & Hughes 1995).
Their ecological and economic importance might
nevertheless be quite considerable, especially for the
medusa phase. Some hydromedusae can occur in high
densities and may consume significant amounts of fish
larvae or compete for the same food resource (e.g.,
van der Veer 1985; Purcell 1986; Matsakis & Conover
1991 ). It may also be that owing to their intense feeding
on primary consumers like copepods, occasional algal
blooms may be promoted by them.

From the viewpoint of evolutionary ecology, hydro¬
zoans also offer other interesting, unsolved questions.
Despite many attempts (cf. Boero etal 1992; Schuchert
1993 ), it is still not known why some species within
the same genus have lost their medusa and others still
spend most of their life cycle in this phase. Most prob¬
ably there will be no single answer for all species, but
the results might offer instructive views on life-history
evolution in marine invertebrates. The present work
hopes to promote such studies by providing the
necessary systematic framework without which
serious ecological studies are not possible (cf. Boero
1984 ). No systematic work can ever be complete and
in this monograph many gaps of knowledge remain.
Wherever possible, the author tried to point out these
gaps and it is hoped this will motivate others to fill
them.

Area of Study

This study reviews all records of marine Antho-
medusae and athecate hydroids from New Zealand,
including the Kermadec Islands and the subantarctic
islands under New Zealand jurisdiction (Fig. 1). This
geo-graphic range includes subtropical to subantarctic
waters. Around the main islands sea temperatures
^how a marked gradient and in summer reach around
2l c C in the far north and 9°C in the far south. Despite
this difference the main islands are rather homo¬
geneous from a biogeographic viewpoint (Morton &
Miller 1968). The Kermadec Islands in contrast are
more tropical and the small islands south of Stewart
Island belong fully to the subantarctic zone. More
information and references on the marine biogeo¬

graphy of New Zealand can be found in, for example,
Ekman (1953), Morton and Miller (1968), Cairns
(1991), Adams (1994).

Terminology

The general morphology of an anthomedusan, an
athecate hydroid, general terminology, and nemato-
cyst types are explained in Figure 2. Following the
functional arguments of Bouillon et al (1988b), the
macrobasic eurytele is here also defined as a eurytele
with a shaft length of more than 2.5 times the capsule
length. Other authors use a minimal shaft length of 4
times capsule length. In the present work, the ratio of
shaft length to capsule length is give as an s value for
all heteronemes.

For more terms see Russell (1953), Naumov (1969),
Millard (1975), and O' Sullivan (1982).

Explanation of technical terms used in this work:

abaxial: position in a medusa facing away from main axis,
abaxial spurs: processes on abaxial side of marginal bulbs
that extend onto exumbrella,
aboral: away from mouth.

adaxial: position or direction in a medusa facing towards
main axis.

adradial: position in a medusa between perradial and
interradial.

apical canal: canal or chamber that originates from the
top of the manubrium and projects into the apical jelly,
apical funnel: funnel-like depression at the apex of a
medusa.

apical: top of a medusa,
atrichous: nematocyst without spines,
autoepizooism: settlement and growth of colonies on
older animals of the same species,
blastostyle: structure carrying gonophores. Not all
blastostyles are homologous organs — in the Euden-
driidae the reduced hydranths bearing gonophores
are also called blastostyles.

capitate: tentacle structure with nematocysts
concentrated in a large, terminal cluster,
caul us: stem-like support of a hydranth.
chordoid: arrangement of gastrodermal cells in a tentacle;

cells are arranged in a single row like a roll of coins,
claviform: club-shaped,
cnidocysts: synonym of nematocysts.
coenosarc: living tissue of a colony,
colony: assemblage of polyps that have a common gastric
system, mostly connected by a system of stolons. The
term colony is frequently used incorrectly by referring
to an upright stem (cormoid), which may be a part of
a colony only (e.g., in colonies that form feather-like
upright stems).

cormoid: erect, branching elements of a colony, e.g., one
single feather-like structure of a Pennaria colony.

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cryptomedusoid: gonophores with no radial canals,
gastrodermis present only as a lamella,
dactylozooid: modified polyps with a defensive function,
comprising tentaculozooids, spiral zooids, etc.
diaphragm: protrusion of gastrodermis partitioning the
gastric cavity of polyps of the Corymorphidae (in
some thecate hydroids also, a perisarcal shelf occurs
near the base of a hydrotheca),
dichotomous: branching pattern in which the youngest
hydranths are distal and branches are forked (see also
monopodial).

distal: position away from centre.

ectodermis: outer germ layer, but often used instead of
epidermis, which should be avoided (see Fautin &
Mariscal 1991).

endodermis: inner germ layer; but often used instead of
gastrodermis, which should be avoided (see Fautin
& Mariscal 1991).

epidermis: outer epithelial layer, derived from ecto¬
dermal germ layer, see also ectodermis.
eumedusoid: gonophore that still has radial canals, but
lacks tentacles; some eumedusoids may be released
and live independently for a short time,
exumbrella: outer part of the bell of a medusa,
fascicled: erect stems composed of several tubes (stems),
filiform: tentacle structure in which nematocysts are
evenly distributed over tentacle,
gastrodermis: inner layer of cells, derived from ento-
dermal germ layer, see also endodermis.
gastrozooid: normal feeding polyps with mouth and
normally with tentacles, without reproductive organs,
gonangium: enclosed reproductive organ composed of
several gonophores.

gonophores: reproductive structures formed during
polyp stage, which may develop into free medusa or
remain fixed, phylogenetically derived from one
medusa bud only. In this work the term is used as in
Rees (1957) or Bouillon (1985a). Some authors
incorrectly use the term in a different sense, applying
it to all sessile repro-ductive structures (here referred
to as sporosacs).

gonozooid: reproductive polyps bearing gonophores,
usually modified gastrozooids that show various
stages of reduction and loss of tentacles,
heterotrichous: nematocyst with spines that vary in size,
holophyletic: monophyletic taxon that includes all
descendants of an ancestral species,
holotrichous: nematocyst with spines along the whole
shaft or thread.

hydranth: actual polyp, consisting of hydranth body,
tentacles, and hypostome.
hydroid: polyp stage of hydrozoans.
hydrophores: perisarcal structures at the base of
hydranths in the family Solanderiidae. Often as two
parallel triangular processes,
hydrorhiza: all structures by which polyps are attached
to the substratum.

hypostome: region around mouth of polyp, mostly

between mouth and first whorl of tentacles,
interradial: position in a medusa between radial canals,
manubrium: feeding organ of medusae, often composed
of stomach and mouth part,
marginal bulbs: tentacle bulbs of hydromedusae,
merotrichous: nematocyst with spines confined to one,
distal, region of its thread.

moniliform: refers to an arrangement of nematocyst
clusters on tentacles; clusters are arranged in bead¬
like rings of tall epidermal cells and in a terminal
knob.

monopodial: branching pattern of stems in which the
oldest hydranth remains at the distal end.
nematocysts: stinging capsules, see Figure 2.
oral: towards mouth.

paraphyletic: group of taxa that does not include all
descendents of one taxon.

parenchymatic: arrangement of gastrodermal cells in a
tentacle, cells are irregularly arranged, no central
lumen present,
pedicel: stem.

peduncle: bulge of the jelly in the centre of the sub¬
umbrella bearing the manubrium, cf. Fig. 27.
perisarc: horny substance covering stolons or stems,
perradial: position in a medusa where radial canals occur,
planula: larva of cnidarians.

plesiomorphy: nonderived character, not suitable for
deducing phylogenetic relationships,
proximal: position closer to centre,
pseudofiliform: tentacles with scattered nematocysts in
a relatively low epidermis along one side and a
concentration of nematocysts in a tall epidermis on
the other side (Petersen 1990).
pseudohydrotheca: covering of hydranth body by a film¬
like, flexible periderm.

spiral zooid: modified polyps without mouth or tentacles
but with a gastral cavity, characteristic of some
Hydractiniidae, typically performing writhing move¬
ments and tending to twist into a spiral,
sporosacs: reduced medusae or gonophores that mostly
remain fixed to the hydranth and release gametes from
there; there is no further implication on its structure
(see also styloid, eumedusoid). See also Millard (1975).
stem: any erect structure bearing hydranths arising from
a hydrorhiza.

stolon: tubular organ at base of polyps by which it
adheres to the substratum.

stolonal: polyps arise from stolons; they may have a
caulus; used here in the sense of colonies without
branching stems. Some authors use this term in a more
restricted sense and apply it to colonies in which
polyps arise from the hydrorhiza without stems
(cauli).

styloid: gonophores without radial canals, without
gastrodermis lamella, and without rudiment of sub¬
umbrella. Some authors (e. g., Werner 1984)
incorrectly synonymise this term with sporosac.
subumbrella: bell cavity of a medusa.

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■■ ^jpomorphy: derived character that is shared by
several taxa and is useful to deduce phylogenetic
relationships.

tenfeculozooids: polyp similar to tentacle in structure,
mostly with a solid core of gastrodermis and no mouth

or gastral cavity. More delicate and slender than spiral
zooids.

vasiform: shaped like a vase, with broad base and slender
top.

velum: iris-like membrane at the bell opening of hydro¬
medusae.

METHODS

Synonyms and References

Sot all synonyms and references to a species are cited
m this study. Only synonyms cited in earlier publi¬
cations dealing with New Zealand species, important
lor the understanding of the scope of the taxon, or
used in the references are given here. Lists of synonyms
can in most cases be found in one of the references
siven. The references given for a species are also not
intended to reflect a complete record. Only references
used for the identification or otherwise important
contributions are given. The thesis of Barnett (1985)
n not an officially recognised systematic publication,
therefore her species names are not synonyms. They
are, however, treated as valid records.

Collection of Material

Live material used for this study was collected mainly
from the Wellington area. Living specimens were also
obtained from near Leigh, Kaikoura, and Portobello
(Dunedin). Hydroids were collected by snorkelling,
SCUBA diving, and once by dredging (Kaikoura).
Depths are usually given as below low-tide mark.

Smaller polyps were mostly detected by collecting
pieces of rock, shells, ascidians, or seaweeds and
examining them in the laboratory under a dissecting
microscope. Where possible, polyps were either
cultivated attached to their original substratum or
growing on 5 ml plastic petri dishes. These were kept
in a shallow plastic container (60 x 40 x 10 cm 3 ) with
running, sand-filtered seawater (3 to 6 litres/min.
flow). The hydroids were fed at least every second
day with living Artemia nauplii or copepods caught
with a plankton net.

Medusae were collected by towing a plankton net
(60 cm diameter opening, 200 pm mesh size) along
jetties for a total length of approximately 300 metres.
Samples were taken regularly (at least every second
week) from jetties near Greta Point and Seatoun
(Wellington Harbour, see Fig. 1). Occasional live
plankton was obtained from Whangateau Harbour
(Leigh) and Portobello. Medusae were cultivated

individually in 5 ml petri dishes or 20 ml bowls at
room temperature or at sea temperature. The water
was changed daily. The medusae were fed every day
(if possible) with Artemia nauplii or pieces of calanoid
copepods. Spawning was induced, where necessary
or possible, by a change from dark (12 h) to light.

Nematocysts were examined using live material
where available. At least ten capsules of each type were
drawn with the help of a camera lucida (CL, tracing
tube). Measurements were then made from the draw¬
ings after the scale had been established by calibration.
Measurements could also be made in increments of
0.5 pm, which is in a sensible relation to the
resolving power of light microscopes. Where possible,
preserved material was also examined. Some material
was cleared by immersion in 50% lactic acid. For the
nomenclature of the capsules see above.

Preserved material of hydroids and medusae were
made available from various institutions (see list
below: Abbreviations). The most significant collection
of Anthomedusae was provided by Mrs Treffery
Barnett (Auckland). These were collected by herself
and others and were the basis of her 1985 thesis.

All new type material was registered and deposited
at the N.Z. Oceanographic Institute of NIWA in
Wellington. Most other newly collected material was
also deposited there, but without individual access
numbers.

Drawings were made whenever possible using
living animals, otherwise formalin-fixed specimens
were used. The bells of preserved medusae can be
distorted and crumpled; in figures depicting preserved
medusae, the umbrella is here shown without crinkles
and corrected for obvious distortions. Such recon¬
structions are normally not difficult where more than
one specimen is available.

Abbreviations

Abbreviations in figures are explained in the captions,
CL indicates that these figures were originally drawn
using a camera lucida. The abbreviation m.w. means
metres of wire (cable length for plankton tows).

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Mafbaur

Westport

WELLINGTON

Island' Te f
Bay Raekaihau

Rangitoto I

Auckland]

Kermadec Islands

■

Chatham Islands

Q. Auckland Island

^Campbell Island

Poor Knights
■Islands

Stewart Island

Fig. 1. Overview maps for study area.

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Abbreviations of institutions:

MCC: Museum of Canterbury, Christchurch, New
Zealand

NIWA: National Institute of Water and Atmospheric
Research, Greta Point, Wellington
NMNZ: National Museum of New Zealand, Wellington

now Museum of New Zealand Te Papa Tonga-
re wa (MoNZ).

NZOI: New Zealand Oceanographic Institute, Well¬
ington (now part of NIWA)

ROM: Royal Ontario Museum, Toronto, Canada
MVM: Museum of Victoria, Melbourne, Australia
ZMC: Zoological Museum Copenhagen, Denmark

LIST OF STATIONS

Stn

Date

Latitude

rs)

Longitude

Depth

(m)

Stn

Date

Latitude Longitude Depth

(*S) (m)

New Zealand Oceanographic Institute Stns

U799

7.8.90

42°33.8' 170°34'

434

X480F

17.10.93

41°20.4’ 179°05.8' 200-400

A439

4.10.58

40°59.6'

174°27.2'

140-160

Z1060

17.8.54

Devonport Wharf, Auckland

A414

28.3.58

23°16'

177°12.5'

surface

ship's hull

B706

13.9.62

41°17.4'

174°41.1'

Z7840

19.2.60

77*42.15 166°19.5'E 351-432

C814

25.2.62

37°40'

178°56.4'

194

Z7863

Mar. 94

Evans Bay, Wellington

C380

28.10.59

38°50'

174°21.5'

Z7864

3.8.94

Goat Island, Leigh

E251

6.4.65

34°35‘

172°35'

Z7865

6.10.94

Te Raekaihau Pt, Wellington

E413

11.10.65

45°12'

171°44'

594

Z7866

26.2.94

Te Raekaihau Pt, Wellington

169

14.5.75

36°11.2'

175°17.7'

Z7867

1.3.94

South Bay, Kaikoura

1357

20.11.77

35°27.8 r

174°44.1'

0-10

Z7868

Feb. 81

Whangateau Harbour

1378

23.11.77

34°09.5 r

172°08.7'

0-24

Z7869

1984

Leigh Marine Reserve

1673

7.9.74

36°26.37'

175°45.60'

Z7870

26.11.93

Doubtful Sound, Fiordland

1674

8.9.74

36°41.85'

175°55.20'

20-30

Z7871

18.1.94

Seatoun, Wellington

J698

10.7.74

37°49.8 r

176°51.8'

6-10

Z7872

7,1.94

Narrow Neck Beach, Auckland 1

1705

11.9.74

37°16'

176°51'

190

Z7873

10.5.94

Aquarium Point, Dunedin

J 899

22.1.76

35°51.0 r

174°28.0'

intertidal

Z7874

1.11.59

Aquarium Point, Dunedin

N356

7.12.74

36°31.3'

175°17.6'

0-50

Z7875

13.3.94

Houghton Bay, Wellington

N371

10.12.74

34°23‘

171°55'

0-200

Z7876

5.9.94

Greta Point, Wellington

0.5

N400

16.12.74

40°26.8 r

175°9.2'

0-53

X401

16.12.74

40-24'

174°52'

100

N404

17.12.74

41°38'

175°19'

0-51

Dana Stns

\413

18.12.74

42°32.5'

173°49.9’

500

-S421

19.12.74

41°24.4'

174°45'

0-100

3624IX

10.12.28

28°19.5' 176°56’ E

100

X433

30.1.75

41°46.1'

171°25.9'

0-25

3626VIII 13.12.28

27°00 177°4T W

1500

X449

1.2.75

44°28.9'

167°38.6’

0-200

3627II

14.12.28

30°08‘ 176°50’ W

4000

N453

2.2.73

46°00.8'

166°36.4'

351

3627VIII 14.12.28

30°08‘ 176°50 1 W

300

X465

5.2.75

47°40.7'

167°01.2'

0-154

3630V

17.12.28

34°24‘ 178°42.5' E

600

Q725

5.3.82

42°25‘

171°06'

363III

18.12.28

35°40 176°40 1 E

1«00

Q726

5.3.82

41°25.50'

172°01.20'

3809

4.9.29

6°14.5' 105°06.5 f E

600

U796

6.8.90

42°34‘

170 o 34.4’

428

3844V

11.10.29

12°05' 96°45' E

1000

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-apical process

manubrium
-exumbrella
gonad

hypos tome

■mesentery

oral tentacles
~f- mouth lip

radial canal

nematocyst pad

: ^\''subumbrella

x \

'circular canal
marginal bulb
—tentacle

perradial adradial

Fig. 2. Anatomical terms used in this study. Top left: hypothetical anthomedusa. Top right: hypothetical polyp stage. Below:
nematocyst types found in this study, all discharged, a) desmonemes, b) atrichous isorhiza, c) basitrichous anisorhiza, d)
heterotrichous anisorhiza, e) microbasic mastigophore, f) microbasic euryteles, g) macrobasic eurytele, shaft longer than 2.5
times capsule length, h) stenotele.

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SYSTEMATICS

Remarks: The arrangement of taxa does not imply a
phylogeny, although for the capitate families a phylo¬
genetic arrangement as proposed by Petersen (1990)
is adopted. Taxa within families are mostly arranged
alphabetically, with new taxa at the end. Within a
species, the more distinctive stage is described first.

SUBCLASS ANTHOATHECATA

Hydrozoans that always have a polyp stage.
Hydranth body not covered by rigid perisarc. Medu¬
sae not colonial, without statocysts, with gonads on
manubrium, with radial canals, with tentacles arising
from bell margin. Cnidome normally includes
desmonemes.

Remarks: Following Cornelius (1992), the name Antho-
athecata is here preferred instead of the compound
name Anthomedusa-Athecata in order to avoid
vestiges of the old dual classification. The Antho-
athecata is a well-defined group that is easy to recog¬
nise. Problems arise only when the group has to be
characterised by cladistic methodology (Wiley 1981).
There is no obvious synapomorphy that solely de¬
limits this group as a holophyletic taxon. Affinities
with the Siphonophora and Laingiomedusae are not
established. All three taxa have gonads on the manu¬
brium as a possible synapomorphy. Desmonemes
occur only in the Siphonophora and Anthoathecata
and may indicate a closer relationship. A taxon
Hydroidomedusae (Bouillon et aL 1992a) which
includes all Hydrozoa except the Siphonophora is
therefore most probably not a natural taxon.

Order FILIFERA

Athecate hydroids with filiform tentacles; only
exceptionally are capitate tentacles present in
tentaculozooids (family Ptilocodiidae). Gonophores
remain either fixed or are released as free medusae.
Manubrium of medusa normally with pronounced
tetraradial symmetry, Cnidome does not include
stenoteles.

Remarks: The absence of stenoteles is an important
synapomorphy that separates the Filifera from the
Capitata (Petersen 1990), Some rare reports of steno¬
teles in otherwise typical filiferan polyps like Bimeria

australis Watson, 1978 require a re-investigation. They
are probably of foreign origin (cf. Calder 1993: 1001)
or a convergently evolved capsule (cf. Bouillon et al.
1986). If real stenoteles should nevertheless be
present inB. australis , then its systematic position must
be re-evaluated thoroughly. The tetraradial symmetry
of the medusa mouth is a plesiomorphy and it is also
present in a few little-derived Capitata. But the
tetraradial symmetry is a useful diagnostic character
to separate most medusae of the Filifera. In this group
the tetraradial symmetry of the manubrium can be
rather pronounced by oral tentacles, oral arms,
perradial lips, or mesenteries.

The Stylasteridae (hydrocorals) are nowadays also
included in the Filifera. They are related to the Hydr-
actiniidae. Because Cairns (1991) recently revised the
Stylasteridae of New Zealand this group is not
included here.

Family CLAVIDAE McCrady, 1859

Solitary or colonial hydroids. Colonies stolonial or
branching, arising from tubular, ramifying hydrorhiza.
Growth in erect colonies monopodial with terminal
hydranths. Perisarc soft or firm, investing hydrorhiza
only or covering both hydrorhiza and hydrocaulus,
occasionally forming cones into which hydranths can
retract. Hydranths elongate, spindle- to club-shaped,
with scattered filiform tentacles. Hypostome conical.
Nematophores present or absent. Gonophores fixed
sporosacs or free medusae, developing from hydro¬
rhiza, hydrocaulus, branches, pedicels, or entire or
reduced hydranths. Medusae bell-shaped. Mouth of
medusa surrounded by four lips, margins of lips with
clusters of nematocysts. With four radial canals and
circular canal. Marginal tentacles solitary, numerous
in adult. Ocelli present. Gonads on interradial walls
of manubrium.

Remarks: The above diagnosis was slightly modified
after Calder (1988) to accommodate Merona and the
new species Rhizogeton conicum which both have the
ability to retract into the perisarc cone enveloping the
caulus.

Until recently, polyps with scattered filiform
tentacles lacking stenoteles could quite securely be
allocated to the family Clavidae, but Boero and
Bouillon (1989) described the polyp of the pandeid
medusa Stomotoca atra . This polyp has scattered

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filiform tentacles and, in the absence of life-cycle
information, would have been classified as a member
of the Clavidae. However, the typical bend-back
behaviour of the closely related genus Amphinema is
also shown by this polyp (Boero & Bouillon 1989, and
also p. 62). This shows on the one hand how important
live observation and complete life-cycle information
are, and on the other that the present system is still far
from satisfactory.

Characteristics of New Zealand genera:
Cordylophora : erect, branching colonies, sessile
gonangia, in brackish water.

Oceania : clavid medusa, without large vacuolated cells
on manubrium base.

Rhizogeton : stolonial colonies with sessile gonophores
arising from stolons.

Turritopsis : erect, branching hydroid colonies, free

medusae, often red or orange colours, medusa with
large vacuolated cells on top of manubrium.

Cordylophora Allman, 1844

Erect, branching hydroid colonies with monopodial
growth and terminal hydranths. Hydranths naked,
spindle-shaped, provided with scattered filiform
tentacles. Gonads in gonangia which grow singly on
cauli of hydranths. The larvae may develop within the
gonangia to a quite advanced stage, sometimes even
young polyps (after Bouillon 1985a).

Type Species: Cordylophora caspia (Pallas, 1771).

Remarks: Only one species is known from New Zea¬
land.

Fig. 3. Cordylophora caspia from life, a) erect colony; scale bar 2 mm. b) nematocysts: microbasic eurytele, same discharged,
desmoneme, same discharged; scale bar 10 fim. c) hydranth with young female gonangium; scale bar 0.4 mm. d) spent
gonangium, same scale as c). e) mature male gonangium, note branching of blastostyle (shaded) and nematocysts at distal
end; scale bar 0.2 mm.

(cc )®®(S

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Cordylophora caspia (Pallas, 1771)

(Fig. 3a-e)

Tubularia caspia Pallas, 1771: 433.

Cordylophora lacustris Allman, 1844: 395.

Cordylophora lacustris: Hamilton 1883: 419.

Cordylophora lacustris var. otagoensis Fyfe, 1929: 813, figs 2-

10 .

Cordylophora lacustris otagoensis: Ralph 1953: 64, fig. 6.
Cordylophora caspia: Naumov 1969: 196, fig. 66.

Material Examined:

Several living colonies from Tomahawk Lagoon, Dunedin,
growing on the underside of stones, collected 14.5.94,
material deposited.

Description: Erect, branching hydroid colonies, up to
30 mm, arising from ramified and anastomosing
stolons. Occasional autoepizooism is present. Erect
stems with monopodial growth, branching several
times irregularly at various angles, ending in up to 10
hydranths. Perisarc thick, smooth, only occasionally
annulated. Hydranths spindle-shaped, up to 1.4 mm
long, with distinct hypostome, which can be nipple¬
shaped. Tentacles filiform, 20-27 in number, scattered
over hydranth body. Tentacles of varying length (up
to 1.4 mm), tapering slightly to three-quarters of basal
diameter, gastrodermis composed of chordoid cells.
Gonads in ellipsoid gonangia arising at the base of
hydranths, covered with perisarc. Initially they possess
a branched blastostyle, which may be reduced to a
simple one later in development. Gametes are released
through distal hole. Male gonangia have a distal cluster
of nematocysts. Nematocysts:

a) microbasic euryteles of very variable size, shaft in
unexploded capsule with conspicuous notch at end
(Fig. 3b), (8-11) x (3.5-5.5) pm, s = 0.9.

b) desmonemes, discharged with 3 coils and bristles
inside of coils, (5-5.5) x (3-3.5) pm.

Colours: Basal parts of stems and branches black,
turning distally into amber; hydranths white; gonangia
opaque.

Tentacle numbers: Median 22, range 19-27, n = 5.
Hydrorhiza diameter: 270-425 pm.

Stems at origin: 320-344 pm.

Records for New Zealand: Esk River, Napier (Hamilton
1883); Tomahawk Lagoon, Dunedin (Fyfe 1929, this
study); Lake Ellesmere, Canterbury (Ralph 1953).

Other Records: Very wide spread in brackish waters,
originated most probably from the Caspian Sea.

Remarks: The height of colonies can reach 100 mm
(Naumov 1969). This species has a high phenotypic
variability which can in part be correlated with
the degree of salinity (Roch 1924; Kinne 1956).

Considering the effect of salinity on morphology,
creation of form names or even subspecies seems
inappropriate.

Fyfe (1929) and Miller (1973) described the develop¬
ment of the reproductive organs. Miller interpreted
the branching structure in the gonads as a blasto style
rather than as a spadix (vestigial medusa manubrium),
making the whole structure a gonangium, which is an
organ composed of several gonophores. This
gonangium corresponds to the one in Corydemirium
(see Fioroni 1977 for discussion of terminology). Eggs
of C. caspia can develop within the gonangia into
planulae (Fyfe 1929).

Oceania Koelliker, 1853

Colonial hydroids with tentacles dispersed over
hydranth body. Clavid medusae with eight or more
marginal tentacles not grouped in clusters. Manu-
briumupona simple, solid, gelatinous peduncle, never
vacuolated.

Type Species: Oceania armata Koelliker, 1853.

Remarks: Only one species is known from New Zea¬
land.

Oceania artnata Koelliker, 1853 (Fig. 4a-c)

Oceania armata Koelliker, 1853: 323; Metschnikoff 1886: 78,
pi. 1, figs 32-39; Mayer 1910:147, figs 80-81; Kramp 1959:
99, fig. 63; Kramp 1961: 65 (cum si/;?.); Kramp 1968: 27,
fig. 67; Brinckmann-Voss, 1970: pi. 4, fig. 2; Bleeker &
van der Spoel 1988: 230, fig. 6; Bouillon 1995: 224.

Material Examined:

Two medusae from NZOI Stn N404, 41°38'S, 175°19'E, 0-
51 m deep, 7-8 mm high, 80 and 120 tentacles, ocelli almost
completely lost.

Description: Medusa stage: Medusa up to 10 mm high,
bell-shaped with flat top, jelly uniformly thin. Manu¬
brium on a shallow peduncle without vacuolated cells.
Manubrium large, up to two-thirds of subumbrellar
height, flask shaped, with large stomach and funnel-
shaped mouth region. Stomach cruciform in cross
section. Mouth rim crenulated, with 4 prominent
perradial lip s. Margin of mouth with a coniinuou s row
of spherical nematocy st clusters. Gonads on interradial
surface of stomach. Four quite broad radial canals,
margins occasionally jagged, ending in circular canal.
Up to 120 tapering marginal tentacles, longer than bell
(preserved sample), gastrodermis chordoid. Each
tentacle with a slight proximal swelling beginning
shortly after origin.

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Fig. 4. Oceania armata. a) mature medusa from preserved
Cook Strait samples; scale bar 1 mm. b) nematocysts of
medusa: heteroneme, desmoneme, microbasic eury tele from
mouth clusters; scale bar 10 |im. c) polyp stage redrawn
from Metschnikoff (1886); note: the polyp stage has not yet
been found in nature; scale bar 0.5 mm.

Origins of tentacles alternately slightly displaced
adaxially and abaxially. With adaxial ocelli.
Nematocysts:

a) heteronemes (microbasic eurytele?), not seen dis¬
charged, (6-7) x (3-3.5) pm.

b) desmonemes, (5-5.5) x (3-3.5) pm.

c) microbasic euryteles from mouth clusters, (8.5-9)
x (3-3.5) pm.

Polyp stage (after Metschnikoff 1886): Hydroid
colonies arising from ramifying stolons. Hydranths on
short, periderm-covered caulus. Hydranth spindle-
shaped with conical hypostome. With around 13
filiform tentacles in up to 4 whorls.

Type Locality: Mediterranean.

Remarks: The polyp of Oceania armata is only known
from the rearing experiments of Metschnikoff (1886)
and has never been found in nature. The origin of
medusae buds is also not known.

Records from New Zealand: Numerous records from
north of East Cape, Kermadec Islands, east of North
Island, Coromandel Peninsula, Tasman Sea (Kramp
1965), Cook Strait (Bouillon 1995).

Distribution: Medusa present in coastal waters in the
tropical and subtropical parts of the Pacific and Indian
Oceans, Mediterranean, Eastern Atlantic from Portu¬
gal to Cap Verde, Azores, West Indies (Kramp 1965).

Turritopsis McCrady, 1857

Hydroids forming erect, branching colonies. In well-
developed colonies base of hydrocauli adnate for some
distance. Hydranths with irregularly scattered filiform
tentacles. Gonophores develop on the hydrocauli and
are liberated as medusae. The medusae are character¬
ised by an apical mass of vacuolated cells on top of
the manubrium (after Bouillon 1985a).

Type Species: Turritopsis nutricula McCrady, 1857.

Remarks: Only T. nutricula is known from New Zea¬
land.

Turritopsis nutricula McCrady, 1857 (Fig. 5a-e)

Turritopsis nutricula McCrady, 1857: 56, pi. 4, figs 1-10, 12-
15, 28a, pi. 5, figs 11, 16-18, 28b; Mayer 1910: 143, figs
10-13, pi. 14-15; Ralph 1953: 64, figs 10 & 18; Russell
1953:115, figs 54A-C, 55, 56, pi. 5, figs 1-5, pi. 29; Kramp

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Fig. 5. Turritopsis nutricula from life, a) adult medusa with developing embryos on the manubrium, shaded area is red-
coloured; scale bar 1 mm. b) nematocysts in pairs of undischarged and discharged capsules, top row from medusa: eurytele
from tentacle, desmoneme, and eurytele from manubrium; lower row from polyp stage: eurytele, desmoneme, and rare
mastigophore; scale bar 10 mm. c) polyp colony with medusa buds; scale bar 2 mm. d) CL-drawing of the stem region
showing the double-layered structure of the perisarc ( ps ), cs\ coenosarc; scale bar 50 mm. e) newly hatched medusa; scale
bar 0.5mm.

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^0)(D®©

1959:100, fig. 64; Kramp 1961: 66 {cum syn.); Kramp 1968:
27, fig. 66; Millard 1975: 76, figs 24F-G; Calder 1988: 8,
figs 5-6 {cum si/;?.); Hirohito 1988: 71, figs 23c-d; Ramil
& Vervoort 1992: 17; Bouillon 1995: 224.

Tubiclnvn rubra Farquhar, 1895: 209, pi. 13, fig. 6 (new
synonym).

Tubiclnvn fructicosa: Hilgendorf 1898: 201, pi. 16, figs 1, la.
Turritopsis pncificn Maas, 1909: 722.

Tubiclnvn rubra: Bale 1924: 228.

Corydendrium zelnndicum Stechow, 1924: 57 (new synonym).
Corydendrium rubra: Stechow, 1924: 58.

[? Not Tubiclnvn rubra: Ralph 1953,]

Material Examined:

Polyps:

Numerous colonies from Wellington Harbour (Queens
Wharf and Evans Bay), reaching up to 5 cm with hundreds
of hydranths, present throughout all seasons, reduced in
summer.

Some living, infertile colonies from Portobello, Dunedin
(May 1994); preserved sample of fertile colony from same
locality, collected 22.11.53 by P. Ralph (in collection of
Marine Laboratory).

Type of Tubiclnvn rubra Farquhar, 1895, Canterbury
Museum, Christchurch, type no. AO 32733, type locality
Wellington Harbour.

Living Mediterranean material from Ischia (Italy), kindly
provided by Dr S. Piraino.

Medusae:

Living adult medusae, found near Ti Point, Leigh (August
1991); Rangitoto Island, Auckland (August 1991, October
1993); Scorching Bay, Wellington (27.1.1994, many). Juven¬
iles of all stages could be found in the plankton of Evans
Bay regularly during spring and summer.

About 10 preserved medusae collected by T. Barnett, 1983-
1984, Leigh Marine Reserve.

Life cycle:

Medusae released from polyp colonies from Evans Bay were
cultivated up to the 16-tentacle stage, when the adult
characters become evident.

Description:

Polyp stage: Hydroids forming initially stoloial, later
erect colonies, arising from attached, ramified stolons.
Branching of stems irregular, reaching 15 mm (up to
50 mm in very large colonies). Branches originate at
acute angles and are adnate for some distance after¬
wards, then curving away and becoming free.
Branches 160-200 pm in diameter, some few expand¬
ing distally; perisarc firm, mostly heavily infested with
detritus and algae, without annulations, terminating
below hydranths. Perisarc double-layered, inner layer
corrugated. Hydranths in material from New Zealand
intensively orange-red. Relaxed hydranths are spindle-
shaped and reach 0.8 mm in height, with 12-20 filiform
tentacles of different length (max. 0.6 mm), highly
contractile, scattered over distal three-quarters of
hydranth; hypostome conical, hydranth not retract¬
able into perisarc, stressed hydranths contract to egg-
shape. Medusae buds arise below hydranths in

perisarc- covered region, mostly one bud, occasionally
more. The medusae buds arise from short stems which
may also be adnate for some distance after their
origin. Also hydranths arising directly from stolons
but belonging to a branching colony can bear medusae
buds. Nematocysts:

a) microbasic euryteles, (7-8) x (3-4) pm, s = 0.9.

b) microbasic mastigophores, rare, in body and cauli
of larger polyps, (5.5-7) x (3-4) pm, s = 0.7.

c) desmonemes, discharged with four coils, (4.5-5.5)
x (2.5-3) pm.

Young medusa: Newly released medusa spherical,
light orange in New Zealand material, manubrium
with 4 perradial lips with nematocyst clusters, manu-
brial peduncle present, base of manubrium and
transition to radial canals with very large vacuolated
gastrodermal cells, 4 interradial orange pads on distal
part of manubrium. With 4 radial canals and circular
canal. With 8 tentacles, arising from bulbs with an
inconspicuous adaxial ocellus. The exumbrella appears
granulated and is slightly opaque. Nematocysts:

a) microbasic euryteles in tentacles, (7-8) x (3-4) pm.

b) microbasic euryteles of manubrium, (8.5-10) x (3-
3.5) pm.

c) desmonemes, discharged with four coils, (5-6.5) x
(2.5-3.5) pm.

Adult medusa: Up to 7 mm high, bell-shaped, higher
than wide, jelly thin, at apex thicker. Stomach cross¬
shaped in axial view, manubrium not longer than bell
cavity. Manubrium with intensively red colour. Four
radial masses of large vacuolated gastrodermal cells
form a compact mass above stomach. Mouth with 4
lips with nematocyst clusters along margin. Four radial
canals which continue through vacuolated gastro¬
dermal masses to manubrium. Gonads on interradial
walls of manubrium, in mature females with develop¬
ing embryos and planulae. Many tentacles (up to 120),
arising from closely spaced tentacle-bulbs, these with
adaxial ocelli. Nematocysts:

a) microbasic euryteles in tentacles, (8-9) x (4-5) pm,
s = 0.9.

b) microbasic euryteles from manubrium, (7-9.5) x (3-
4) pm, s = 0.7.

c) desmonemes of tentacles, (5.5-7) x (3.5-4.5) pm.

Type Locality: Charleston Harbour, South Carolina,
USA.

Remarks: Turritopsis nutricula is a cosmopolitan species
and frequently found in intertidal (polyp) and coastal
waters (medusa). The medusae, even young ones, are
readily recognised by their vacuolated gastrodermal
cells at the base of their manubrium. Additionally, the

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size and red colour of the adult make the medusa con¬
spicuous and they can even be discerned from small
boats. Also young medusae are intensively orange.
Colouring, however, may depend on food sources.

The polyp was found all year round in Wellington
Harbour, often in rather dense populations and often
with mussels. The characteristic double-layered
structure of the perisarc of the cauli (Fig. 5d) makes
them identifiable even when the hydranth is not well
preserved (Ramil & Vervoort 1992). This structure is,
however, not everywhere as obvious as given in the
figure.

The examined material from New Zealand agrees
well with descriptions provided by other authors (see
synonymy). Only the presence of microbasic mastigo-
phores in the polyp stage has not been reported so
far. These nematocysts occur in the hydranth body and
cauli, sometimes in a cluster of up to 20 capsules. They
do not occur in the tentacles and it may be that not all
polyps have them, especially young polyps. The
present material from New Zealand was compared to
living T. nutricula hydroids from the Mediterranean
(Ischia, Italy). These specimens likewise do have the
mastigophores. There are nevertheless some differ¬
ences between these populations. Ihe Mediterranean
polyps were much lighter coloured and showed a
broader branching. The newly hatched medusa was
quite colourless and its vacuolated cells much less
developed. Their tentacles showed a unique trembling
behaviour not seen in medusae from New Zealand.
While most of these characters can be explained by
differences in the environment, a closer examination
of the life cycle of the Mediterranean population seems
appropriate to test whether they are a different species
as recently postulated by Calder (1988).

After examination of the type specimens of Tubi-
clava rubra Farquhar, 1895 it became evident that this
sample belongs to Turritopsis nutricula. Although frag¬
mented, the type material of Tubiclava rubra is well
preserved. The sample consists of pieces of macro¬
algae (presumably holdfasts of Macrocystis pyrifera as
given in the original publication) covered with
hydroids, mostly by a clavid type. They form erect
colonies that are branched 2-3 times. The cauli are
tubular, not dilating distally, branching at acute angles
with the new branch being adnate to the other for some
distance. The hydranths are typical clavids with up to
16 scattered filiform tentacles. The perisarc shows the
characteristic double-layered structure. Few gono-
phores are present, arising below hydranths in
perisarc-covered region, up to 3 in one row along the
caulus. They arise on short stems, which are also
adnate for some distance. The gonophores are obvious
incipient free medusae as the manubrium, 4 radial
canals and the tentacles are visible. It is not possible

to distinguish Farquhar's sample from Tur-ritopsis
nutricula . The general abundance of this species at the
type locality also argues in favour of this interpretation.
Stechow (1924) has already noted that Tubiclava cannot
be the correct genus for Farquhar's hydroid, as
Tubiclava is not branching, if the genus is valid at all.
Stechow's proposed name Corydendrium rubra there¬
fore becomes a synonym of Turritopsis nutricula.
Similarly, Stechow (1924) and Bale (1924) proposed
that Hilgendorf s (1898) Tubiclava fructicosa from
Dunedin should better be referred to T. rubra.
Stechow's proposed new name of Corydendrium zelan-
dicum is therefore another synonym of Turritopsis
nutricula. Tubiclava rubra depicted by Ralph (1953, fig.
5A) cannot belong to the genus Tubiclava because it
has branched hydrocauli. Her illustration shows a
quite robust hydroid (from Christchurch ?) which may
be not Turritopsis nu triad a, as she described it as having
fixed gonophores on separate stems. This description
was, however, copied from Farquhar (1895). There¬
fore, it remains unclear what material she had in hand.

Records from New Zealand: Hydroid: Wellington
Harbour (Ralph 1953; this study); Portobello, Dunedin
(Ralph 1953; this study). Medusa: Bare Island, Hawke
Bay (Kramp 1928, as T. pacifica); Cook Strait (Kaberry
1937); Auckland (Kramp 1965; Jillett 1971; this study);
Leigh (Barnett 1985; this study); Wellington Harbour
(this study).

Distribution: Circumglobal in warm to temperate
waters, e.g.. Northeast Pacific (Fraser 1948), South
Africa (Millard 1975), North Atlantic (Russell 1953),
Pacific (Kramp 1965), Papua New Guinea (Bouillon
1980).

Rhizogeton Agassiz, 1862

Stolonal hydroid colonies, either sessile or with a
perisarc-covered caulus; hydranths with scattered
filiform tentacles. Nematophores absent. Gonophores
arise from hydrorhiza and are sessile sporosacs.

Type Species: Rhizogeton fusiforme L. Agassiz, 1862.

Remarks: The diagnosis as given by Bouillon (1985a)
was modified to accommodate Rhizogeton sterreri
(Calder, 1988) and Rhizogeton conicum n. sp. The
emendation now also allows the presence of perisarc-
covered cauli. Mainly based on shape of the hypo-
stome, Calder (1988) split the genus as defined
above into two genera ( Rhizogeton and Rhizodendrium).
Because the gonophores of the type species for Rhizo¬
dendrium are not yet known, Calder's proposal is not

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followed here.

Two species are known from New Zealand with
the following distinguishing characters:

Rhizogeton conicum : base of hydranth with conical
perisarc tube into which hydranth can retract.
Rhizogeton sp.: hydranths sessile, with very short peri¬
sarc collar at hydranth base; with elongated vege¬
tative propagules.

Rhizogeton conicum n. sp. (Fig. 6a-d)

Material Examined:

1 colony from western side of Te Raekaihau, 30.1.1994,
subtidal, on sponge.

1 hydranth from western side of Te Raekaihau, 26.2.1994,
subtidal, on barnacle, infertile.

1 colony from western side of Te Raekaihau, 13.3.1994, 1-
2 m, on sponge, fertile, part of it used for slide preparation,
deposited as holotype H-652, remainder as paratype P-
1082.

1 colony from Houghton Bay, Wellington, 13 m, on ascidian,
infertile, hydranths up to 4 mm, deposited.

Description: Hydroid colonies arising from sparingly
ramified, loosely adhering stolons. Stolons covered
with perisarc. Hydranths on unbranched cauli.
Perisarc smooth or wrinkled, thin and filmy on caulus
of hydranth where it forms an inverted cone up to

2 mm long into which hydranths can partially or
completely retract; size of cone is variable. Hydranths
cylindrical, 1-4 mm long and 0.3 mm diameter, with
up to 20 filiform tentacles. Tentacles organised in distal

3 to 4 whorls with 4 tentacles in each whorl, addition¬
ally few more irregularly scattered tentacles below.
Tentacle gastrodermis chordoid. Hypostome dome¬
shaped. Gonophores sessile sporosacs, arising directly
from stolons, spherical, with larger filmy perisarc, up
to 4 eggs embedded in tissue with many nematocysts.
Nematocysts:

a) microbasic euryteles of polyp, shaft projecting at
an angle of 45° or less, (10-14) x (4-5.5) pm, s = 0.8.

b) microbasic euryteles of gonophores, more bean¬
shaped than other eurytele, shaft discharges side¬
ways, (11-13) x (4.5-5) pm, s = 0.7.

c) desmonemes, (5-7) x (3-4) pm.

Colour: white and transparent, gonophore white and
opaque.

Fig. 6. Rhizogeton conicum n. sp., from life, a) incipient gonophore (left) arising from stolon, fully expanded (middle) and
contracted (right) hydranth; scale bar 0.5 mm. b) mature gonophore enclosed in thin, flexible perisarc, same scale as a), c)
female gonophore tissue treated with lactic acid to make it transparent, CL; scale bar 0.2 mm. d) nematocysts in pairs of
native and discharged capsule: euryteles of polyp, desmonemes, euryteles of gonophore; scale bar 10 pm.

0®g)(

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Type Iadcality: West of Te Raekaihau, Wellington.

Remarks: Rhizogeton conicum is somewhat unique
within its genus as it has a conical perisarc covering
of the caulus into which the polyp can withdraw
completely, or occasionally in older polyps at least
partially (the cone may not always be large enough to
accommodate the whole polyp). Within the genus
Rhizogeton the polyps of R. conicum are similar to those
of R. sterreri (Calder, 1988). Unfortunately the gono-
phores of the latter are not known, but R. conicum is
distinct from R. sterreri by the longer perisarc cone and
its ability to retract into it. Rhizogeton sterreri is not
able to do so (D. Calder, pers. comm.). Other similar
species are Rhizogeton ezoense Yamada, 1964 from Japan
and R. fusiforme Agassiz, 1862. But both of them have
markedly oblong gonophores. The polyps of R. conicum
also resemble polyps of the genera Merona or the little
known Tubiclava. There is indeed some potential to
mistake R. conicum for a Merona species, because
occasionally a few folliculinid ciliates are attached to
the stolons. Although smaller, the thecae of these
ciliates have a deceptively similar morphology like a
perisarc covering of a nematophore as found in Merona
(cf. Millard 1975). An examination of the soft tissues
with a compound microscope, however, reveals the
difference.

No males have been found so far, although the
species seems to occur quite regularly at its type
locality. The living tissue of the female gonophore
was very opaque and internal structures could be
made visible only by clearing them with lactic acid.

Records from New Zealand: Known only from Well¬
ington's south coast.

Rhizogeton sp. (Fig. 7 a-b)

Material Examined:

Several colonies (1-2 cm 2 each) on oysters collected inter-
tidally near Portobello Marine Laboratory, 10.5.1994,
colonies transferred and cultivated on plastic petri dishes
for three months, no gonophores developed, vegetative
propagules developed abundantly and colonies spread
to other objects.

2 colonies on mussel and sponge collected 28.6.1994 near
Greta Point, Wellington, 0.5 m, infertile.

Description: Hydroid colonies with hydranths arising
directly from attached, ramified stolons. Stolons
covered by perisarc. Hydranths up to 4 mm high, with
shallow basal collar of perisarc, conical hypostome
and up to 20 filiform, tapering tentacles scattered in
distal half of polyp body. With small red pigment
granules in epidermis of hydranth body and tentacle
bases.

Fig, 7. Rhizogeton sp. from life, a) part of polyp colony with
two propagules growing from stolons (right); scale bar 0.5
mm. b) nematocysts: desmoneme, microbasic eurytele
discharged, same undischarged; scale bar 10 ^im.

Spherical to elongated (2 mm long) structures with
a perisarc-covered caulus can develop from stolons.
These structures (propagules) can detach and are able
to re-attach at other sites where they first turn into
stolons and later into new colonies. Nematocysts:

a) microbasic euryteles, (7.5-8) x (3-3.5) pm, s = 1.

b) desmonemes, (5.5-6) x (3-3.5) |Jm.

Remarks: Because no information on gonophores could
be obtained, this species was not given a specific name.
The hydranths are very similar to those of Rhizogeton
nudum Broch, 1909 (cf. Millard 1975; Jones 1992, but
these authors disagree on gonophore morphology).
The propagules observed in the present study are not
known for R. nudum. These propagules or their
younger stages can easily be mistaken for incipient
gonophores. Several of them from various colonies
were checked thoroughly with the help of a compound
microscope, but gametes or similar cells were never
detected. The propagules are hollow and have the
same basic organisation as a hydranth body. They are

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very sticky and re-attach rather easily. After a short
time, the cultivated colony had spread to various
points in the aquarium, although the individual
colonies remained quite small. The species was rather
abundant near Portobello and was also found by P.
Ralph around 1955 (unpublished notes kept by Porto¬
bello Marine Laboratory).

Records from New Zealand: Portobello; Wellington
(this study).

Family CALYCOPSIDAE Bigelow, 1913

Hydroids, where known, living in the prebranchial
cavities of ascidians. Polyps colonial, cauli not branch¬
ing, arising directly from hydrorhiza. Hydrorhiza
formed as a plate. Polyps with up to five irregular
whorls of filiform tentacles. Gonophores arise from
polyps and are liberated as free medusae. Medusae
without apical projection and without gastric ped¬
uncle. Mouth with four simple or crenulated lips.
Gonads on manubrium, simple or folded. Four or eight
radial canals, simple or branching. Blindly ending
centripetal canals may be present. Marginal tentacles
hollow, four, eight, or more in number. Tentacles lack
basal bulbs or basal swellings but have terminal
swellings with nematocysts. There may also be small
or rudimentary tentacles. The basal portion of the
tentacle is often adnate to the exumbrella. Ocelli only
exceptionally present (Bouillon 1985a).

Remarks: Characteristics of genera known from New
Zealand:

Bythotiara: medusa with four primary radial canals that
may branch distally, no centripetal canals.
Calycopsis : medusa with blindly ending centripetal
canals.

Bythotiara Guenther, 1903

Hydroids as given in family diagnosis. Medusae as in
family diagnosis but with four radial canals which may
branch, without centripetal canals. Gonads interradial,
with transverse furrows. With or without secondary
tentacles.

Type Species: Bythotiara murrayi Guenther, 1903

Remarks: Pages et al (1992) provided a table with the
characteristics of all known Bythotiara medusae.

Characteristics of species known from New Zealand:
Bythotiara murrayi’. medusa up to 20 mm, radial canals
branching.

Bythotiara parasitica : hydroids living in ascidians.
Bythotiara sp.: medusa up to 4 mm, radial canals not
branching.

Bythotiara murrayi Guenther, 1903 (Fig. 8)

Bythotiara murrayi Guenther, 1903: 424, pi. 10, figs 4-5;
Russell 1940: 515; Russell 1953: 215, pl.13, fig. 1, text-
figs 113a-b,114a-b, 115-116; Kramp 1959:125, figs 1,132;
Kramp 1961: 118; Kramp 1968: 54, fig. 142; van der Spoel
& Bleeker 1988: 167, fig. 17; Pages et al 1992: 7, fig. 7.

Material Examined:

1 medusa from Dana Stn 3627V1I1, 30°08'S, 176°50'W,
14.12.1928,300 m wire, identified by P. Kramp as Bythotiara
murrayi, very damaged.

1 medusa from Dana Stn, 3844V, 12°05'S, 96°45'E, 11.10.1929,
1000 m wire, identified by P. Kramp as B. murrayi, very
damaged, not recognisable.

Description (after Kramp 1968): Medusa up to 20 mm
high, with thick walls; stomach small, with 4 inter¬
radial gonads with transverse furrows; radial canals
generally 4, bifurcate (but additional branching may
occur); long tentacles as many as ends of radial canals;
some small secondary tentacles and minute tentacles.
Long tentacles ending in terminal swellings. Nemato¬
cysts (after Russell 1940):

a) ? euryteles, (17-20) x (10-11) pm.

b) desmonemes, on tentacle tips, discharged with 5
coils, (13-14) x (6) pm.

Fig. 8. Bythotiara murrayi , modified after Guenther (1903)
and other sources; no scale given, rear tentacles and radial
canals not shown for reasons of clarity.

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Type Locality: 52°18.1'N / 15°53.9'W (SW of Ireland).

Records from New Zealand: Near Kcrmadec Islands
(Kramp 1965).

Distribution: Eastern parts of Atlantic from Norway
to South Africa; Mediterranean; tropical parts of the
Indian Ocean; Papua New Guinea (Kramp 1968; Pages
et al. 1992).

Bythotiara parasitica (Kirk, 1915) n. comb.

(Fig. 9 a-d)

Ascidioclava parasitica Kirk, 1915: 146, pi. 1, figs 1-6.
Endocrypta liuntsmani: Trebilcock 1928: 1; Ralph 1953: 66,
fig. 4.

[Not Endocrypta liuntsmani Fraser, 1911]

? Endocrypta parasitica : Briggs &: Gardiner 1931: 186.

Maierial Examined:

1 colony found in 1 out of 10 Pyura rugata Brewin, 1948 (cf.
Millar 1982) collected in Scorching Bay,Wellington Har¬
bour, 3 m, 13.9.1994, medusae buds present.

5 colonies found in 5 out of 6 Pyura rugata collected in Eve
Bay (outside Wellington Harbour), 4-7 m deep, 10.10.1994,
medusae buds present, but only the two most advanced
were liberated and could be cultivated for a few days only,
polyp material deposited.

Description:

Polyp stage: Hydroids living in ascidians, especially
Pyura rugata. Colonies loosely attached to the feather¬
like buccal tentacles and also around their base. Polyps
with non-branching cauli, arising directly from a plate-
like hydrorhiza which is not covered by perisarc.
Polyps up to 3 mm high. Larger, relaxed polyps with
a cylindrical caulus, egg-shaped body, and a conical
to spherical hypostome (depending on state of con¬
traction). On distal part of hydranth body 3 (max. 4)
whorls of filiform tentacles, total number 20-30, up
to 1 mm long, all of similar length, rather thick. Caulus
can be as long as body, transparent, with large gastro-
dermal cells, not covered by perisarc.

Gonophores (1-4) arise at the basal end of the
hydranth body where it turns into the caulus. Gono¬
phores not covered by perisarc, and liberated as
medusae. Nematocysts:

a) microbasic euryteles, (10-13) x (4-5) pm, s = 0.9.

b) desmonemes, discharged with 3 coils, (6-9.5) x (3-
5) pm.

c) atrichous isorhizas, rare, (13-16) x (6-8) pm.
Colours: hydranth body pink to purple, opaque.

Newly released medusa: Bell-shaped, 1 mm high,
higher than wide. Jelly moderately thick. Exumbrella

covered with nematocysts which are lost during the
following development. Dilated velum spanning onc-
third of radius. Manubrium somewhat less than half
of subumbrellar height, cruciform in cross-section,
mouth simple, cruciform. Four radial canals and ring
canal present. Tentacle bulbs absent. With 4 perradial
tentacles, these shorter than bell height, ending in an
intensively orange-coloured terminal swelling. No
ocelli present. Nematocysts:

a) microbasic mastigophores, some almost like
euryteles, only onexumbrella, (18-20) x (17-19) pm,
s = 0.7.

b) microbasic euryteles with long shaft, very faint
distal swelling, (7.5-10) x (4-5.5) pm, s = 1.7.

c) microbasic euryteles with short shaft, (6-8.5) x (4-
6) pm, s =1.

d) desmonemes, undischarged thread with ropy
texture, discharged with 5 to 6 coils, (10-14) x (4-
6.5) mm.

Type Locality: Wellington Harbour.

Remarks: Bythotiara parasitica very much resembles B.
liuntsmani (Fraser, 1911), but according to Brinckmann-
Voss (1979) and Rees (1979b), B. liuntsmani medusae
do not have desmonemes and they are released at a
size of 2-2.5 mm, whereas B. parasitica medusae are
released at a size of 1 mm. Therefore, they are seen
here as different species. Also the medusa of B. stilbosa
Mills & Rees, 1979 does not have desmonemes.
Bythothiara stilbosa can, furthermore, be distinguished
by the possession of macrobasic euryteles. Briggs and
Gardiner (1931) reported B. parasitica (as Endocrypta
parasitica) from the Great Barrier Reef in Australia.
They give no information on nematocysts and there¬
fore this record has to be verified.

Brinckmann- Voss (1979) and Rees (1979b) found
R liuntsmani in various ascidian species. During the
present study, many dozens of various solitary ascid¬
ians were examined from different localities around
Wellington, but infected animals could be found only
outside or near the harbour entrance and in waters
below 3 m deep. Infected animals were confined to
one species, identified with some hesitation as Pyura
rugata , a very abundant ascidian aTOund Wellington.
Kirk (1915) also reported a rare occurrence in Pyura
pachydermatina (syn. Boltenia pachydermatina ). As for
some Hydractinia species (Buss & Yund 1989), it should
be tested whether the various ascidian species host
not different but morphologically indistinguishable
species.

As experienced by Brinckmann-Voss (1979) with
Bythotiara liuntsmani , B. parasitica polyps can be
cultivated only for a short time outside their host.
Polyps from nature had their stomachs filled with uni-

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Fig. 9. Bythotiara parasitica, from life, a) polyp colony with medusae buds, the two polyps to the left are contracted; scale
bar 0.2 mm. b) nematocysts of polyp phase in pairs of native and discharged capsules: microbasic euryteles, desmonemes,
atrichous isorhiza; scale bar 10 pm. c) newly released medusa, same scale as a), d) nematocysts of newly released medusa:
microbasic mastigophores (top), microbasic euryteles with long shaft, microbasic euryteles with short shaft, desmonemes;
same scale as b). e) Bythotiara sp., male, most probably the adult of B. parasitica, cultivated from medusae taken from the
plankton near Seatoun; scale bar 1 mm.

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@(D®©

cellular algae, predominantly diatoms. But in culture
they occasionally also fed on Artemia nauplii which
were brought close to their mouth. Medusae released
from the polyps could not be cultivated for more than
a few days.

The polyp colonies collected near the Wellington
Harbour entrance released medusae which were
identical to ones regularly found in the plankton of
the harbour and which could be grown to maturity.
These medusae were very likely B. parasitica. But as
long as the medusae of B. parasitica have not been
cultivated to maturity, it seems more correct to des¬
cribe these medusae separately as Bythotiara sp. (see
below).

Records from New Zealand: Polyps known only from
Wellington (Kirk 1915, and this study).

Oiher Records: ?Great Barrier Reef, Australia (Briggs
& Gardiner 1931).

Bythotiara sp. (Fig. 9e)

Material Examined:

Fairly regularly present in plankton taken at Seatoun (Well¬
ington Harbour entrance), dates and numbers (in
brackets): 8.12.93 (1), 13.1.94 (5), 14.4.94 (4), 18.7.94 (10),
29.8.94 (2), 30.8.94 (5), 11.10.94 (4). All were infertile,
ranging in size from 1 mm (morphology as in fig. 9c) to a
size of 2.5 mm. The medusae from July could be cultivated
to maturity. They were fed on Artemia and parts of fish
larvae and kept at room temperature. Some material
deposited.

1 juvenile medusa collected 15.5.1994 in plankton near
Portobello Marine Laboratory (Dunedin).

Description: Mature medusae 3-3.8 mm high, bell¬
shaped umbrella with thick apical jelly (thickness
one-quarter of height), lateral jelly moderately thick.
Dilated velum spanning one-third of radius. In
younger stages there may be some nematocysts on
exumbrella. Manubrium half as long as subumbrellar
height, cruciform in cross-section, with 4 simple
perradial lips. Mouth margin provided with many
tightly set nematocyst clusters and long cilia. Gonads
on upper half of manubrium, only slightly separated
in perradial position, with an interradial furrow and
in males with 2-3 horizontal folds. These folds are
quite variable from animal to animal and can be absent
in females. Four simple, smooth radial canals and a
circular canal are present. No tentacle bulbs present.
Perradial tentacles 4, without basal swelling. Basal part
adnate to exumbrella for some distance. Tentacles
shorter than bell height, terminating in an intensively
orange-coloured knob with nematocysts. No ocelli

present. Nematocysts:

a) microbasic mastigophores, present on exumbrella
of younger animals, (14-18) x (11-15) pm, s = 0.8.

b) microbasic euryteles with long shaft, very faint
distal swelling, (7-9) x (3-4) pm, s = 2.

c) microbasic euryteles with short shaft, on manu¬
brium, (6.5-10) x (4-7.5) pm, s = 0.9.

d) desmonemes, on tentacle tips, undischarged thread
with ropy texture, discharged with 5 to 6 coils, (9.5-
13) x (4-6) pm.

Remarks: The orange tentacle tips make this medusa
rather conspicuous and easy to recognise. As indicated
above, this medusa is almost certainly the adult
medusa of B. parasitica. The medusae were grown from
specimens which looked like young ones of B. para¬
sitica and their morphology did not change much. Also
their nematocysts are identical. However, it seems
more correct to wait for a successful rearing experi¬
ment before it is definitely allocated to B. parasitica.
The distinction from the very similar Bythotiara
huntsmani is again possible through the presence of
desmonemes and the smaller size of the bell.

Young stages of this medusa are very difficult to
cultivate. Only when they become older are they easier
to feed. In contrast to B. huntsmani , however, they seem
not very temperature-sensitive as they could be
maintained at room temperature. Medusae show an
interesting feeding reaction — upon contact with prey
the tentacles bend into the subumbrella, which closes
by contracting not only the velum but the margin.

Records from New Zealand: Cook Strait (Bouillon 1995,
as ?B. huntsmani); Wellington and Portobello (this
study).

Other Records: Not known outside New Zealand.

Calycopsis Fewkes, 1882

Calycopsid medusae with mainly four unbranched
radial canals and with four or more centripetal canals
arising from the ring canal, blindly ending or joining
the cruciform base of the stomach. Gonads trans¬
versely folded, frequently forming eight adradial rows
of deep transverse furrows. Basal portion of tentacles
adnate to umbrella margin; all tentacles hollow,
nematocysts only in the terminal knob (Kramp 1961).

Type Species: Calycopsis typa Fewkes, 1882.

Remarks: An overview of the genus is given in Kramp
(1959). No polyp stages are known. At present there
is only one species known from New Zealand waters.

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Cahjcopsis bigelozui Vanhoeffen, 1911 (Fig. lOa-b)

Cahjcopsis bigelowi Vanhoeffen 1911: 218, fig. 12; Kramp
1957a: 21, map fig. 4; Kramp 1959: 127, fig. 136; Kramp
1961: 119; Kramp 1968: 56, fig. 149; van der Spoel &
Bleeker 1988: 167, fig. 16.

Material Examined:

1 medusa, 16 mm high, from NZOI Stn X480F, 41°20.45,
179°05.8'E (east of Cape Palliser), 17.10.1993, 200-400 m.
For comparison, 17 medusae from Dana Stn 3809 (near
Sumatra & Java, 6*14.55,105°06.5'E), 4.9.1929, 600 m wire,
up to ~1 cm in size, det. P. Kramp.

Description: Mature medusa up to 16 mm high, dia¬
meter 16 mm, spherical. Jelly thick (3 mm), soft and
adhesive. Velum somewhat sunken into bell. Manu¬
brium length two-thirds of bell cavity, with cross¬
shaped base. Mouth with slightly undulating margin
and with 4 small perradial lips. With 4 large interradial
gonads leaving only perradial parts of manubrium
visible. Gonads with 8 adradial rows of up to 16 hori¬
zontal folds. With 4 complete radial canals, those near
manubrium dilated and thus forming mesenteries.
With 4 additional centripetal canals which almost
reach manubrium base but end blindly, terminal
region somewhat broadened. All radial canals are
rather broad and are connected to the equally broad
circular canal. There are no tentacular bulbs present.
With 4 perradial, 4 interradial, and 8 adradial tentacles.
Perradial and interradial tentacles are all fully
developed, most of the adradial tentacles are also fully
developed but some are shorter and developing. In
addition to these 16 long tentacles there are 16 very
short tentacles, one in between each pair of longer
tentacles. All tentacles are hollow and at their base
adnate to the exumbrella. All full-length tentacles end
in a terminal swelling with a distal concentration of
nematocysts. Tentacles shorter than full-length ones
have no distal swelling, but a concentration of
nematocysts is present distally. No ocelli present.
Nematocysts:

a) desmonemes, on tentacle tips, discharged with 5
coils, thread with spiral pattern of small bristles,
(11-12) x (5-6) pm.

b) heteronemes, rare, not seen discharged (14-
15) x (7-8) pm.

Type Locality: Gulf of Aden.

Remarks: Cahjcopsis medusae are quite rare and are
known only from a few individuals from few locations
(Kramp 1959). Therefore little is known about their
morphological variation and development. The single
medusa collected from New Zealand is rather well
preserved, but differs somewhat from the existing
descriptions and also the examined specimens from

Fig. 10. Cahjcopsis bigeloiv, from Cape Palliser. a) medusa,
only frontal tentacles shown; scale bar 5 mm. b) nematocysts:
desmoneme, same discharged, rare heteroneme; scale bar
li pm.

Indonesia. The major difference was in the bell size
and morphology of the gonads. While the other medu¬
sae had fewer (around 10) and looser gonad folds, the
medusa from New Zealand had up to 16 folds which
were very tightly set. They resembled the gonads of
the other large Cahjcopsis medusae. The other differ¬
ence concerned the tentacles. While other C. bigelowi
are known with 8 fully grown tentacles (Kramp 1968;
van der Spoel & Bleeker 1988), the present samples
also had most of the adradial tentacles fully grown.
However, a few of them were not fully grown and
some did not yet have the terminal swelling. The very

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short tentacles were again as described by others.
These differences are not seen as sufficient to create a
new species, and the medusa from New Zealand
appears to be an older, more developed specimen. The
examined material from Indonesia had gonads with
visible eggs. It may therefore be that C. bigelowi
continues its growth after gonads have become
mature. Such a post-mature growth is known in
other medusae and has been the cause of some taxo¬
nomic problems (cf. Kubota 1992).

Another species of calycopsid medusa with four
blindly ending centripetal canals is Calycopsis borch-
grevinki (Browne, 1910). This species most probably
also occurs in southern parts of New Zealand due to
its circumpolar occurrence in subantarctic waters.
However, this species has a very different gonad struc¬
ture, with the gonads sunken into the manubrium with
many circular openings (cf. Browne 1910; Moore 1984;
van der Spoel 1992). Kramp (1968, fig. 146) copied the
somewhat atypical figure of Vanhoeffen (1911).

Records from New Zealand: Cape Palliser (this study).

Other Records: Gulf of Aden; Cape of Good Hope;
Indo-Malay an region; tropical Indian Ocean; in deeper
waters down to 600 m (Kramp 1965; van der Spoel &
Bleeker 1988).

Family BOUGAINVILLIIDAE Luetken, 1850

Stolonal or branching hydroid colonies, exceptionally
sessile. Branches covered with perisarc. The perisarc
terminates either at the base of the hydranths or con¬
tinues over them as a thin, filmy pseudohydrotheca.
The polyps have one or more distal whorls of filiform
tentacles. The gonophores are either fixed sporosacs
or are liberated as medusae. Gonophores develop
mostly from cauli or branches, occasionally from
hydrorhizae or rarely from modified hydranths.
Medusae have a simple circular mouth with oral
tentacles inserted above it. Oral tentacles are either
simple or branched. Gonads differentiate either in
interradial, adradial, or perradial positions, or encircle
the manubrium. Four radial canals and a circular canal
are always present. Marginal tentacles are simple,
either two, four, or more solitary tentacles, or groups
of tentacles arising from four, eight or sixteen bulbs.
With or without ocelli.

Remarks: The Bougainvilliidae are an assemblage of
quite different hydroids and difficult to delimit from
other families (Calder 1988). Future revisions will
certainly be quite extensive. Therefore, the more usual
classification of Bouillon (1985a) is used here and not

that of Calder (1988). Bougainvilliid polyps without
reproductive structures are often not distinguishable
from those of the family Pandeidae.

Characteristics of genera known from New Zealand:
Bougainvillia : free medusae with four groups of
marginal tentacles.

Dicoryne : dimorphic polyps, gonophores develop on
gonozooids as sporosacs that are liberated.
Gravelya: monomorphic polyps with gonophores
reduced to sporosacs arising also from stolons.
Koellikerina : free medusae with 8 groups of marginal
tentacles.

Bougainvillia Lesson, 1836

Colonial hydroids, branching or not. Hydranths may
be covered by thin, wrinkled perisarc called pseudo¬
hydrotheca. Hydranths with one whorl of filiform
tentacles, which are never enveloped by perisarc. The
gonophores arc liberated as free medusae. Medusae
with four oral tentacles which may be branched or
not, attached above level of mouth. Marginal tentacles
all alike and set into four perradial groups. Normally
with ocelli. The gonads are either in perradial,
interradial, or adradial positions (after Bouillon 1985a).

Type Species: Bougainvillia macloviana (Lesson, 1830).

Remarks: With the exception of B. inaequalis, most
Bougainvillia polyp colonies are not identifiable alone
and information on the adult medusa is needed. For a
review of Bougainvillia species see Vannucci and Rees
(1961).

Characteristics of species present in New Zealand:

B. aurantiaca: medusa small, up to 4 tentacles per bulb,
without ocelli.

B. fulva: medusa less than 14 mm high, 15-20 short
tentacles per bulb, with 8 adradial gonads, well
separated interradially.

B. inaequalis : polyp colonies with wrinkled perisarc
throughout, medusa unknown.

B. macloviana : medusa up to 15 mm high, with large
peduncle, bulbs with 30-60 tentacles.

B. muscus : small medusa, up to 3 mm, with not more
than 5 tentacles per bulb.

B. platygaster : medusa up to 12 mm high, with flat and
quadrangular stomach, up to 10-13 tentacles per
bulb, crescent-shaped ocelli, oral tentacles with
very short trunk.

B. vervoorti : medusa up to 7 mm, up to 30 tentacles
per bulb, large cross-shaped base of stomach
bearing gonads.

B. dimorpha: medusa up to 5 mm, not more than 10
tentacles per bulb, sexually dimorphic, male with
gonads.

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Bougainvillia aurantiaca Bouillon, 1980 (Fig. 11)

Bougairtvillia aurantiaca Bouillon, 1980: 309, fig. 1; Goy et al.
1991: 107, fig. 19.

Maierial Examined:

2 medusae, 1.2-1.5 mm, collected by Treffery Barnett in June
to July 1984, Leigh Marine Reserve.

Description: Medusa with bell-shaped umbrella, up
to 1.5 mm, with slightly thicker jelly at apex.
Manubrium simple, conical, length half to two-thirds
of bell cavity, with or without a very slight peduncle
(preserved material), with simple mouth and 4
perradial oral tentacles. Oral tentacles branching 2 to

3 times and ending in nematocyst clusters. Basal trunk
of each oraltentacle very long, up to half of total length.
Gonads as 4 separated elongated pads in interradial
position on manubrium, may contact each other in
perradial position. Radial canals and circular canal

Fig. 11. Bougainvillia aurantiaca from Leigh; scale bar
0.2 mm.

rather narrow. Four perradial tentacle bulbs, each
bearing 2-3 tentacles. Ocelli absent. Nematocysts:

a) heteronemes (microbasic euryteles ?), (6.5-7) x (3.5-
4) (im.

b) desmonemes (5.5-6) x (3.5-4) pm.

Polyp stage: unknown.

Type Locality: Laing Island, Papua New Guinea.

Remarks: The medusa may reach 1.9 mm in height
(Bouillon 1980) or even 2.5 mm (Barnett 1985). The ten¬
tacle bulbs are intensively orange in living animals
(Bouillon 1980).

The absence of ocelli and the long basal trunks of
the oral tentacles make this species rather distinctive.
A similar species is Bougainvillia muscoides which also
has no ocelli, but can be two times as large and has
short basal trunks. Bouillon (1995) reported this species
for New Zealand, although with a query. His material
was re-examined for this study (NZOI collection, Stn
N449). All other Bougainvillia medusae from this
collection have lost their ocelli because of fixation. It
is therefore impossible to identify the small Bougain¬
villia species because no information on the ocelli can
be obtained. It seems better therefore to disregard the
record of B. muscoides from New Zealand.

Records from New Zealand: Leigh Marine Reserve
(Barnett 1985).

Other Records: Papua New Guinea (Bouillon 1980),
Mediterranean (Goy et al. 1991).

Bougainvillia fulva Agassiz & Mayer, 1899 (Fig. 12)

Bougainvillia fulva Agassiz & Mayer, 1899: 162, pi. 2, fig. 6;
Maas 1905: 10, pi. 1, fig. 8, pi. 2, figs 9-10; Vanhoeffen
1911: 207; Kramp 1928: 47, figs 21-23; Kramp 1961: 77;
Kramp 1968: 33, fig. 84; Bouillon 1995: 223.

Material Examined:

3 medusae from NZOI Stn N421, 41°24.4'S, 174°45'E, 0-
100 m, 19.12.1974.

Description: Medusa bell-shaped, up to 11 mm, higher
than wide. Jelly thick (one-fifth bell height), bell margin
lobed through 4 perradial furrows. Manubrium broad,
cruciform in cross section, length about one-quarter
to one-third of bell cavity, with 4 perradial oral
tentacles. Each oral tentacle branched up to 5 times
and all endings with a nematocyst cluster. Basal trunks
of oral tentacles very short. Gonads as 8 oblong
adradial pads on manubrium wall, well separated
interradially and perradially. Four moderately broad
radials and a ring canal. Four broad, V-shaped per-

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)®@©

Fig. 12. Bougainvillia fulva; scale bar 2 mm.

radial tentacle bulbs with up to 10 short tentacles.
Polyp stage unknown.

Type Locality: Fiji Islands.

Remarks: Ocelli occur at tentacle bases and are slightly
elongated (Vannucci & Rees 1961); the stomach may
bud medusae (Kramp 1965) and the eggs are not
covered by a layer of nematocysts (Hartlaub 1909b).

The bell margin of New Zealand Bougainvillia fulva
was found to be lobed due to perradial notches in the
bell margin which allow the tentacles more freedom
to move. Vanhoeffen (1911) also described the same
morphology. The existing descriptions and figures of
B. fulva are not satisfying and some are even contra¬
dictory (e.g., Agassiz & Mayer 1902, and Bigelow 1909
on jelly). Bougainvillia fulva as described by Uchida
(1927: 221, fig. 40) may not be this species because its
gonads are interradial.

Records from New Zealand: Off Christchurch (Kramp
1965); Cook Strait (Bouillon 1995).

Distribution: Tropical parts of the Indian Ocean and
eastern Pacific Ocean (Kramp 1968; map in van der
Spoel & Bleeker 1988).

Bougainvillia cf. inaequalis Fraser, 1944

(Fig. 13a-b)

Bougainvillia inaequalis Fraser 1944: 51, pi. 5, fig. 20; Ralph
1953: 63, fig. 3; Vannucci & Rees 1961: 67, table 4.

Material Examined:

1 colony on shell from NZOI Stn E251, 34°35 , S, 172°35 , E,
(North Cape), 6.4.1965, 9 m, dredged on sand, infertile,
not fascicled, only tentatively assigned to this species.

Description (after Vannucci & Rees 1961): Erect,
branching colonies. The stem and sometimes the main
branches fascicled. Stems 7 mm high and straight,
there are only small branches or pedicels given off
singly or in clusters. The perisarc is very much
wrinkled and extends up to the base of tentacles on
the hydranth. The latter has from 8-10 tentacles.
Medusa buds are borne singly or in clusters on the
pedicels of the hydranths. Medusa unknown.

Type Locality: Louisiana, USA.

Remarks: As long as its life cycle remains unknown,
Bougainvillia inaequalis is an insufficiently described

Fig. 13. Bougainvillia cf. inaequalis from Stn E251. a) ter¬
minal portion of colony showing wrinkling of perisarc, CL;
scale bar 0.2 mm. b) nematocysts: microbasic eurytele, same
discharged, desmoneme; scale bar 10 pm.

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species and should not be used for identifications
outside its type locality. Its main characteristic is the
strong wrinkling of the perisarc, which in other species,
however, is always to some extent dependent on the
environment. While the figures given by Ralph (1953)
agree well with Fraser's (1944) description, the sample
examined in the present study (Fig. 13a) differs in being
non-fascicled. Only the strong wrinkling of the peri¬
sarc was used to allocate it tentatively to this species.
More observations from the type locality and New
Zealand are needed to resolve the doubtful status of
this species.

Records from New Zealand: Lyttleton Harbour (Ralph
1953); ?North Cape (this study).

Other Records: Louisiana, USA.

Bougainvillia macloviana (Lesson, 1830)

(Fig. 14a-c)

Cyanea macloviana Lesson, 1830: 118, pi. 14, figs 3D-D".
Hippocrene macloviana : Haeckel 1879: 90, pi. 5, figs 1-2.
Hippocrene : Benham 1909: 306, pi. 12, figs 1-2.

Perigonimus maclovianus : Vanhoeffen 1910: 284, fig. lOa-c.
Bougainvillia macloviana Mayer 1910:160;Hartlaub 1911: 156,
fig. 139; Russell 1953:173, figs 86-88B (cum si///.); Kramp
1959: 109, fig. 85; Kramp 1961: 78; Kramp 1968: 32, fig.
78; Millard 1975: 96, fig. 33A-C; O'Sullivan 1982: 38, fig.
16, map 15; Pages etal. 1992: 3, fig. 1; Bouillon 1995: 223.

Material Examined:

About 6 preserved medusae collected by K. Westerkov
(obtained through T. Barnett), Perseverance Harbour,
Campbell Island, March 1984, size range 8-11 mm high
with up to 60 tentacles per bulb.

2 medusae from NZOI Stn N453, 46°00.8'S, 166°36.4'E, off
Puysegur Point, Fiordland.

Description:

Medusa: Diameter 8 -11 mm (maximally up to 15 mm),
bell as high as wide or slightly higher than wide, with
quadrangular margin, with 4 broad interradial longi¬
tudinal furrows, jelly moderately thick, thickened at
apex. Stomach short, on a well-developed cone-shaped
peduncle. Four narrow perradial lobes of stomach
extend along peduncle. Manubrium with 4 perradial
oral tentacles, not extending beyond umbrella margin,
each branching 5-7 times, all endings with a nemato-
cyst cluster. Basal trunks of oral tentacles rather short.
Gonads on perradial lobes of manubrium that extend
along peduncle. With fairly narrow radial canals and
ring canal. Four crescent- or V-shaped perradial mar¬
ginal bulbs, about half as wide as interradial space,
with 30-60 solid tentacles on each tentacle bulb,
arranged in a double row. Adaxial ocelli correspond¬
ing in number to the tentacles present on bulbs at base

Fig. 14. Bougainvillia macloviana. a) mature medusa from
Campbell Island, scale bar 1 mm. b) nematocysts of medusa:
microbasic eurytele and desmoneme, scale bar 10 Jim. c)
polyp stage redrawn from Vanhoeffen (1910); note: the polyp
stage is not yet known from New Zealand; scale bar 0.5 mm.

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0 ®®(

of each tentacle, shape round, colour red. Eggs covered
with euryteles. Nematocysts:

a) microbasic euryteles, on tentacles and eggs, (7-
8) x (3-4) mm, s = 0.8.

b) desmonemes, (6-7) x (4-4.5) mm.

Polyp stage (after Vanhoeffen 1910, not known from
NZ): Stolonial hydroid colonies, infrequently branch¬
ed, cauli several millimetres high, but often low-lying.
Perisarc smooth or wrinkled, but not annulated, con¬
tinued as thin pseudohydrotheca over hydranth.
Hydranth small, with up to 16 tentacles. Medusa buds
reach 1 mm in diameter and originate from stolons or
stems. Medusae may be released with more than 2
tentacles per bulb (not in South Africa, see Millard
1975).

Type Locality: Falkland Islands.

Remarks: B. macloviana is restricted to cold, subantarctic
waters. There it may occur in rather high numbers. It
also occurs in the North Atlantic, where it has most
probably been transported on ship hulls.

Records from New Zealand (medusa): Auckland and
Campbell Island (Benham 1909; Roberts 1972), south
tip of South Island (Bouillon 1995).

Other Records: Falkland Island, Kerguelen, North Sea
(Russell 1953), South Africa (Millard 1975), Mozam¬
bique, Benguela Current (Pages et cil 1992).

Bougainvillia muscus (Allman, 1863) (Fig. 15a-e)

Eudendrium ramosum : van Beneden, 1844: 56, pi. 4.

[not Eudendrium ramosum (Linnaeus, 1758)]

Perigonymus muscus Allman, 1863: 12.

Bougainvillia ramosa: Russell 1938a: 152; Ralph 1953: 63, fig.

9, not fig. 25; Russell 1953: 153, pi. 8, fig. 1, pi. 9, figs 4-

5, text-figs 74A-C; Kramp 1959:109, fig. 91; Kramp 1961:

81 (cum syn.); Vannucci & Rees 1961: 53; Kramp 1968:

34, fig. 87; Millard 1975: 97, figs 33E-H; Hirohito 1988:

97, figs 34b-f.

Bougainvillia muscus-. Calder 1988: 24, figs 19-20 (cum syn.).

Material Examined:

Several living polyp colonies, up to 50 mm high, fascicled,
from Mahanga Bay, Wellington Harbour, 4 m, collected
30.3.94, medusae released and cultivated to maturity, some
material deposited.

Adult living medusa, from plankton from Evans Bay, 18-
27.4.1994, 7 specimens, 1-2.5 mm, up to 4 marginal ten¬
tacles, oral tentacles once- or twice-branched, some
material deposited.

Preserved colony, 70 mm, fascicled, fertile, from Doubtful
Sound, collected 26.11.1993, 7 m, only tentatively assigned

to B. muscus .

Preserved colonies from fouling panels, collected by S.
Turner from Waitemata Harbour, Auckland, 4 m, fas¬
cicled, up to 30 mm high, most probably this species.
Approx. 5 preserved medusae collected by T. Barnett, Leigh
Marine Reserve, 1984,

Description:

Polyp stage: Hy droids forming erect, branching col¬
onies reaching 50 mm in height, arising from attached,
ramified stolons. Stem fascicled (depending on age),
branching profusely and irregularly. Branches forming
a pointed angle with stem. Perisarc in colonies from
nature infested with detritus, may be wrinkled but not
regularly annulated. Perisarc may continue as thin film
over body of hydranth as a pseudohydrotheca. Stolon-
isation (search stolons) common in all parts of colony,
especially in cultivated material. Hydranths with
conical hypostome and 1 whorl of about 12 tentacles.
Medusa buds arise singly or in groups from hydranth
cauli and smaller branches, smaller than hydranths,
with short stalks, conical when young. Nematocysts:

a) microbasic euryteles, (7-8) x (4-4.5) mm, s = 0.8.

b) desmonemes, discharged with 4 coils, (4-5) x (2.5-
3) mm.

Young medusa: Medusa after release approximately
0.6 mm high, bell-shaped, apical canal may be present.
Tubular manubrium with 4 very short, unbranched
oral tentacles. With 4 marginal bulbs, each with a pair
of slightly beaded tentacles. Marginal bulbs with 2
adaxial red ocelli. There are no exumbrellar nemato¬
cysts present. Nematocysts:

a) microbasic euryteles, (5.5-8) x (2.5-4) mm, s = 0.7.

b) desmonemes, (4-5.5) x (2.5-3) mm.

Mature medusa: Up to 2.5 mm, bell as broad as high,
jelly slightly thicker apically. Bulbous manubrium,
length approximately half of subumbrellar height. Oral
tentacles once- or twice-branching. With 4 marginal
bulbs, each with up to 4 tentacles and ocelli. Ocelli
round. Gonads interradial but reaching perradius,
bulging. Mature eggs are covered with a layer of
euryteles. Nematocysts:

a) microbasic euryteles, (5.5-7) x (3-4) mm.

b) desmonemes, (4-4.5) x (2.5-3) mm.

Type Locality: Torquay, Devon, Great Britain.

Remarks on Variation: Bougainvillia muscus is a very
variable species. From the literature (e.g., Russell 1953;
Vannucci & Rees 1961; Edwards 1966b) the medusa
can reach 4 mm in size with up to 9 tentacles per bulb,
and the oral tentacles may branch up to 4 times.
However, mature animals of only 1 mm size can also
be found. More than 5 tentacles per bulb are rare

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(Vannucci & Rees 1961). Barnett (1985) reported sizes
up to 3.5 mm and 9 tentacles per bulb. Only a few of
Barnett's Bougainvillia medusae could be re-examined.
While the mature animals having fewer than 5
tentacles per bulb are doubtless B. muscus, the ones
having up to 9 are other species (some of them have
crescent-shaped ocelli). Unfortunately not enough
material and information became available to identify
and describe them properly. Younger females of B.

dimorpha n.sp. may be also be mistaken for B. muscus .
Therefore, especially for New Zealand material,
Bougainvillia medusae with more than 5 tentacles per
bulb most probably do not belong to B. muscus.

In the present life-cycle study, maturity was attained
even at 0.8 mm (cultivated animal) with 2 tentacles
per bulb only. Adults from the plankton in Wellington
were found to be not larger than 2.5 mm, 4 tentacles
per bulb maximally.

Fig. 15. Bougainvillia muscus , all from life, a) colony from Wellington Harbour; scale bar 5 mm. b) terminal branch of a
colony with medusa buds which has been kept in culture for about four days. Compared to samples directly from nature,
cultivated colonies tend to have larger hydranths without a pseudohydrotheca, scale bar 0.5 mm. c) newly released medusa;
scale bar 0.2 mm. d) adult male medusa grown from hydroid shown above, 8 days old, scale bar 0.2 mm. e) nematocysts of
polyp or medusa: microbasic eurytele, same discharged, desmoneme, same discharged; scale bar 10 (im.

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cc)(l

General Remarks: Bougainvillia muscus is better known
under its synonym B. ramosa, but Calder (1988) showed
that only the former name is valid. Although for a
correct identification the medusa must be grown to
maturity, large, fascicled Bougainvillia colonies origi
nating from harbours where the medusa is known can
quite reliably be referred to B. muscus (in New Zea¬
land). One sample of large, fascicled Bougainvillia was
obtained from Doubtful Sound. It differed from the
other samples in lacking a detritus cover. Because the
mature medusa is not known and also almost no
information on medusae from Fiordland is available,
the identification is very tentative.

The medusa depicted by Ralph (1953, fig. 25) is not
B. muscus , but B. britannica copied from Mayer (1910).
Ralph admitted, however, that the medusa was not
known from New Zealand.

Bougainvillia muscus hydroids are very easy to culti¬
vate and will produce many medusae, which are also
easy to cultivate. Especially in culture, but also in
natural samples, stolons (search stolons) often issue
from terminal branches. By this, detached parts of the
colony can rapidly fix themselves to any substratum.
The search stolons also help to colonise neighbouring
erect structures like other hydrozoans. Bougainvillia
muscus can spread very rapidly in an aquarium.

Records from New Zealand: Polyp known from
Portobello; West Coast; Coromandel (Ralph 1953);
Wellington Harbour; Waitemata Harbour (Auckland);
?Doubtful Sound (this study). The medusa is known
from near Goat Island, Leigh (Barnett 1985) and
Wellington Harbour (this study).

Other Records: East Australia (the hydroid in Aus¬
tralia, Amboina, China and Japan). Northwestern
Europe, Mediterranean, Black Sea, West Africa, New
England, Brazil (all after Kramp 1968), Papua New
Guinea (Bouillon 1980, medusa), ?South Africa (Mil¬
lard 1975).

Bougainvillia platygaster (Haeckel, 1879) (Fig. 16)

Hjppocrene platygaster Haeckel, 1879: 91.

Bougainvillia platygaster Kramp 1957: 9, text-fig. 1, pi. 3, figs
1-6; Kramp 1959: 109, fig. 89; Kramp 1961: 80; Kramp
1968: 34, fig. 86; Bleeker & van der Spoel 1988: 230, figs
89; Pages et al. 1992: 4, fig. 2; Bouillon 1995: 227, fig.l.

Material Examined:

1 medusa, NZOI Stn N404, 41 0 38’S, 175°19 , E, 0-51 m,
17.12.1974,7 mm high, well-preserved but no ocelli visible.

Description (after present material and Kramp 1957):
Medusa with globular umbrella 4 -7 mm (up to 12 mm)
high, with slightly flattened apex, jelly very thick, sub-

Fig. 16. Bougainvilla platygaster from Cook Strait; scale
bar 1 mm.

umbrella cylindrical, bell margin with perradial
notches. No peduncle present Manubrium very broad
but very short and quadrangular, with 4 perradial
oral tentacles. Oral tentacles arising close to radial
canals and branching dichotomously up to 6 times
immediately from their point of origin, no basal trunk
present. Oral tentacles end in nematocyst clusters.
Medusae may be budded from manubrium or from
polypoid structures arising from the corners of the
manubrium. Gonads interradial pads, flat, not
divided interradially. With 4 small but broad tri¬
angular marginal bulbs with 10-13 short tentacles.
With adaxial ocelli, crescent-shaped with concavity
turned outwards. Polyp stage unknown.

Type Locality: Island of Trinidade (Brazil) (Neotype
after Kramp 1957).

Remarks: The bell margin of Bougainvillia platygaster
was lobed due to clefts in the perradial position allow¬
ing the tentacles to move more freely (see also p. 29).

Records from New Zealand: Cook Strait (Bouillon 1995).

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Other Records: Tropical parts of the Atlantic and Bougainvillia vervoorti Bouillon, 1995

Indian Oceans, Malayan Archipelago, Bismarck Sea, (Pig. 17a-e, 18a-e)

Fiji Islands, Benguela Current (Bouillon 1995).

Bougainvillia vervoorti Bouillon, 1995: 228, figs 2, 3a.

Fig. 17. Bougainvillia vervoorti, from life, a) adult male medusa, rear tentacles not shown; scale bar 1 mm. b) manubrium
and gonads of a female medusa, oral tentacles removed; scale bar 0.5 mm. c) frequently found juvenile parasitic Narcomedusa;
scale bar 0.5 mm. d) tentacle bulbs of adult, CL; scale bar 0.2 mm. e) juvenile medusa from the plankton; scale bar 1 mm. f)
nematocysts; microbasic eurytele, desmoneme; scale bar 10 Jim.

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^0)(D®©

Material Examined:

Holotype H-619, NZOI Stn B706, 41°17.4' S, 174*47.1 1 E /
(end of Clyde Quay, Wellington Harbour), 9 m, female,
collected 13.9.62, kept in NIWA type collection.

Paratypes P-970, same locality, 20.9.62, 2 damaged speci¬
mens; P-971, same locality, collected 28.9.62, 4 specimens.

Ca. 50. living adult medusae, plankton Evans Bay and Sea-
toun, Wellington Harbour, collected November to Decem¬
ber 1993, some deposited.

1 male medusa collected at Portobello, Dunedin, 13.5.94,

Polyp colonies grown from planulae produced by medusae,
cultivated for more than 4 months, material deposited.

Polyp colony from Evans Bay, growing on tube of serpulid
polychaete, collected 30.11.93, 1 m; colonies released
medusae which were cultivated for 16 days until specific

characters became evident (see Appendix), material
deposited.

Description:

Medusa: Adult medusa bell-shaped, 4-10 mm high
and slightly smaller diameter. Umbrella with no or
shallow interradial furrows. Jelly thick, at apex one-
third or more of bell height; a slight peduncle may be
present; basal part of bell quadrangular, bell margins
between bulbs concave. Manubrium with large, per-
radial extension of base, heights of extensions increase
up to the middle then taper. Manubrium length one-
quarter of bell cavity, with 4 long oral tentacles,
branching 5-7 times, ending in nematocyst knobs.

Fig. 18. Bougaiiwillia vervoorti, from life, a) polyp stage with medusae buds found on tubes of serpulid polychaete, scale
bar 0.3 mm. b) newly liberated medusa; scale bar 0.5 mm. c) 8-day old medusa; same scale as b). d) 16-day old medusa,
gonads are visible but not fully developed; scale bar 1 mm. e) nematocysts: microbasic eurytele, desmoneme; scale bar 10 pm.

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branching not always dichotomous. Gonads in 4 pairs
on adradial sides of basal extensions of manubrium,
not contacting each other perradially, in contact inter-
radially, bulging slightly towards interradial. Eggs
covered with nematocysts. Basal manubrial extensions
continue as 4 radial canals. With 4 tentacle bulbs facing
downward, with U-shaped epidermis, each with 18-
30 long and flexible tentacles and corre-sponding
number of ocelli. Ocelli adaxial, dark-red, round, on
tentacle bulb near base of tentacle.

Nematocysts:

a) microbasic euryteles, shaft shorter than capsule,
leaving approx. 45° to main axis, (5-7.5) x (2.5-
3) pm.

b) microbasic euryteles of eggs, (6.5-8) x (3-4) pm.

c) desmonemes, (5.5-6.5) x (3-4) pm.

Other observations: Egg size 160 pm, including
nematocyst layer. Colours: specimens not fully mature
have an intensively emerald green manubrium.

Polyp and young medusa: Hydroid colonies arising
from attached, ramified stolons. Polyps mostly
stolonal, cauli rarely branched once, perisarc covering
of cauli slightly wrinkled, may extend as thin, adher¬
ing film over hydranth body. This pseudohydrotheca
may be absent in older polyps. Hydranths with conical
hypostome and one whorl of 8-10 (up to 12) filiform
tentacles of unequal lengths, projecting alternately up-
and downwards. Medusae buds arise on short stems
either from cauli or stolons. Young medusa oval, jelly
evenly thick, base quadrangular, apical canal present,
manubrium cylindrical with no basal extensions, with
4 unbranched oral tentacles with terminal capitae, 4
thin radial canals, 4 marginal bulbs each with 2
tentacles and 2 ocelli, these dark brown-red, adaxial.
Growth stages, see Appendix 6.1. Nematocysts of
polyps:

a) microbasic euryteles, (6.4-8) x (2.5-3.5) pm.

b) desmonemes, (4-5) x (2.5-3) pm.

Hydranth length: 500 pm (culture), 270-400 pm (fertile
colonies from nature); hydranth diameter 100 pm;
caulus length 0.36-1.2 mm (culture), up to 2 mm
(nature); stolon diameter 60-124 pm; caulus diameter
80-100 pm; tentacle length up to 0.6 mm.

Type Locality: Clyde Quay, Wellington Harbour, 9 m.

Remarks on Life-Cycle Observations: The life cycle of
B. vervoorti was observed in two ways. Adult male
and female medusae from the plankton (they spawned
after reaching the size of 4.2 mm and 16 tentacles per
bulb) were kept in finger bowls at room temperature.
The resulting eggs developed into planulae which
swam for more than one week without meta¬
morphosis. Metamorphosis was then induced by

adding a small piece of Corallina alga (5 times the size
of a planula). Three planulae could be followed as they
settled and metamorphosed on the fragment. The
primary polyps fed vigorously on parts of Artemia and
grew stolons to an underlying microscope slide. The
colony was kept in running filtered seawater and fed
daily. After 28 days post-metamorphosis, the colony
had spread to a diameter of 20 mm and the first
medusa bud on a caulus was observed. Later, more
developed on stolons also. No branching of cauli was
observed.

A colony of Bougainvillia growing on a tube of a
serpulid polychaete collected from Evans Bay was
cultivated in running seawater. The colony was mostly
stolonal: stems were only rarely branched once. The
polyps initially fed well on Artemia and many medusae
buds developed on stolons and cauli (Fig. 18a) while
the hydranths became increasingly smaller. After some
time they were unable to feed on Artemia (reproductive
exhaustion). The released medusae (Fig. 18b) were
cultivated at room temperature with daily feeding. The
development of a representative animal is given in
Appendix 6.1. After 16 days the medusae had devel¬
oped to a stage where it was evident that it was B.
vervoorti (Fig. 18d, Appendix 6.1). The maximal size
was 3.6 mm with 10 marginal tentacles per bulb. The
gonads had just started to develop.

Some size differences were found between the
polyps from nature and those grown from medusae
gametes. Such differences can be explained by different
degrees of reproductive exhaustion.

General Remarks: Bougainvillia vervoorti medusae seem
to be seasonal in Wellington Harbour. Their main
occurrence in this study, and also in earlier samples,
was during spring time. They were not found after
January. Up to 80% of the B, vervoorti medusae were
parasitised by several young narcomedusae (Cunina
sp.. Fig. 17c). The narcomedusae have an elongated
manubrium which they insert into the mouth of B.
vervoorti. Apparently they parasitise on the stomach
content. The narcomedusae attach themselves to the
subumbrella with their tentacles. In such parasitised
B. vervoorti , the gonad development is severely
retarded or inhibited, but the bell size reaches com¬
parable sizes as in non-parasitised ones.

Although they are very different, by following a
formal diagnosis as given by Kramp (1968), B. vervoorti
can be confused with B. fulva Agassiz & Mayer, 1899.
Bougainvillia fulva (p. 20) differs from B. vervoorti in
having only half as many tentacles per bulb, short
tentacles, ocelli on the tentacle base and not the bulb,
lacking large basal extension of the stomach, and
lacking nematocysts on its eggs.

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Records from New Zealand (medusa): Wellington
Harbour (Bouillon 1995 and this study), near Kaikoura
(Bouillon 1995), near Cape Foulwind (Bouillon 1995),
Portobello (this study). Polyp known only from Well¬
ington Harbour.

Other Records: Not known outside New Zealand.

Bougainvillia dimorpha n. sp. (Fig. 19a-h)

Material Examined:

Ca. 100 medusae collected in Evans Bay, February-April
1994. One male medusa; collected 7.3.94 deposited as holo-
type H-638, several males and females as paratype P-1065.

3 crosses made between typical male and female forms, all
3 resulting in planulae which settled and metamorphosed;
ten released medusae of this colony (second generation)
were cultivated, but only 2 females reached maturity.

Description:

Medusa stage: Adult medusa bell-shaped, normally
3-4 mm high, exceptionally up to 6 mm high, about
as broad as high, side walls only slightly curved, apical
jelly up to one-third of height, shallow peduncle may
be present, otherwise top of subumbrella flat, bell base
quadrangular, dilated velum spanning approximately
half of radius. Manubrium distally cone-shaped,
shorter than one-third of bell height, less in males, with

4 mostly dichotomously branching capitate oral ten¬
tacles, each with about 8 capitae. Stomach base with 4
perradial, laterally compressed extensions. In females
extension short, bearing on each side the 8 gonads.
Female gonads are in contact in interradial position-
eggs are covered by a layer of euryteles and arranged
mostly in one layer in the gonads. Male gonads
typically well separated from centre, located on lateral
sides of basal extensions of manubrium. Basal exten¬
sions of male manubrium a narrow tube at origin then
increasing in height where gonads are located. With
4 radial canals and circular canal, all narrow. With
triangular- to heart-shaped marginal bulbs, these much
smaller than space in-between them, each bulb with
7-10 tentacles and a corresponding number of round,
dark-red ocelli. Ocelli on bulbs. Tentacles expanded
longer than twice bell size. Nematocysts:

a) microbasic euryteles of marginal and oral tentacles,
(7-8) x (3.2-4) mm, s = 0.8.

b) microbasic euryteles of eggs, (6.5-7) x (2.5-3) mm.

c) desmonemes of tentacles, with 4 coils, (5-5.5) x (3-
3.5) mm.

Polyp stage and young medusa: Hydroid colonies
arising from attached, ramified stolons. Cauli only
occasionally branching once, reaching a height of up
to 3 mm. Perisarc can form a pseudohydrotheca over
hydranth body. Hydrocauli up to 2 mm. Hydranths

spindle-shaped, about 0.4 mm long, with a conical
hypostome, with 6-10 filiform tentacles in one whorl
but pointing alternatingly up- and downwards, length
about 0.5 mm. Medusae buds arise at right angles
from stolons and from cauli, up to 3 per caulus, bud
diameter up to 0.2 mm. Newly released medusa bell¬
shaped, 0.8 mm high, diameter 0.6 mm; evenly thin
jelly, bell base quadrangular, very slight interradial
furrows present; no apical canal present, with 2 ten¬
tacles and 2 red ocelli per marginal bulb; manubrium
short, oral tentacles unbranched and very short.
Nematocysts of polyps and additional measurements:

a) microbasic euryteles, (7-8) x (3-4) |im, s = 0.8.

b) desmonemes, (3.6-4) x (2.5-3) |im.

Stolon diameter 40-50 pm; caulus diameter 40-60 pm.

Variations: One male medusa with a manubrium and
gonad morphology of normal females was observed.
The length of the basal extensions of the gonads in
females can also vary between individuals by approxi¬
mately a factor of two.

Type Locality: Greta Point, Wellington Harbour.

Live-Cycle Observations: Three pairs of adult medusae
showing the typically different morphologies of the
gonads were used for crossing experiments. All three
crosses resulted in many larvae. The planulae were
induced to settle on very small (1 square mm) flakes
of oyster shell. Settlement and metamorphosis was
very rapid and could be followed. The polyps were
fed daily and grew very rapidly. The colony remained
mostly stolonal and cauli branched only occasionally
(Fig. 19h). Medusae buds appeared after 18 days, from
both stolons and cauli. The released medusae were
quite difficult to cultivate and most did not attain
maturity. Only two females reached maturity (one
spawned). They reached 2.6-3 mm in height in 19 days,
had oral tentacles three times branched, and more than
5 marginal tentacles per bulb. At the time of spawning
all marginal tentacles had been lost. The gonads were
on a cruciform extension of the manubrium base. They
were within the range of morphologies observed in
nature.

Remarks: Male and female medusae of Bougainvillia
dimorpha were first believed to be separate species.
However, their co-occurrence at high densities in
Evans Bay over a short time period indicated a di¬
morphic species. This was then shown by the crossing
experiments which resulted in good fertilisation
success and many larvae. The cultivation of second-
generation medusae was difficult. More observations
are needed to demonstrate the development of the
male gonads.

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Females of B. dimorpha are quite similar to those of
B, muscus . But, in the material from Wellington, they
were quite distinct as B. muscus never had more than
4 tentacles per bulb and were much smaller. Young
stages of B. vemoorti are also similar to B. dimorpha.

But the latter has fewer tentacles (< 10) at maturity,
whereas B. vervoorti with 10 tentacles are not yet
mature. Most males of B. dimorpha are quite distinct
with their gonads separated from the manubrium. Ihe
male medusa bears some resemblance to B. pyramidatn

Fig. 19. Bougainvillia dimorpha n. sp., from life, a) Adult male medusa, very similar to type specimen; scale bar 1 mm. b)
manubrium of a female medusa of similar size as animal in a); scale bar 0.2 mm. c) to f) aboral and side view of a gonads of a
typical male (c, d) and female (e, f) medusa of the same size (4 mm, seven tentacles per bulb); scale bar 0 4 mm. g) Nematocyst
of polyp and medusa, in pairs of intact and discharged capsules: microbasic euryteles, desmonemes; scale bar 10 |im. h) Part of
polyp colony, branching and with medusae buds, grown from crosses of typically dimorphic medusae; scale bar 0.2 mm.

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(cf. Edwards 1964, figs 3-4). In contrast to the latter,
B. dimorpha medusae have no or a much smaller
peduncle. The polyp stage also very different, B. pyra-
midata having a large, fascicled colony (Edwards 1964).

Records from New Zealand: Known only from Evans
Bay, Wellington.

Dicorijne Allman, 1859

Polyps polymorphic, forming erect colonies, branching
or not, covered by perisarc which terminates on
hydranth body or its base. Hydranths with one whorl
of filiform tentacles which are never enveloped by
perisarc. Gonophores are produced on modified
hydranths (gonozooids) and are liberated as ciliated
sporosacs that swim freely (after Bouillon 1985a).

Type Species: Dicoryne conferta (Alder, 1856).

Dicoiyne conybearei (Allman, 1864) (Fig. 20a-d)

Heterocorriyle conybearei Allman, 1864: 365.

Dicoryne conybearei: Ashworth & Ritchie 1915: 257, fig. 3,
pis 6-8, figs 1-15; Hirohito 1988: 99, fig. 35; Cornelius et
nl. 1990: 124, fig. 4.7.

Material Examined:

1 female and 1 male colony, Greta Point, 1 m, on gastropod
shell, 15.12.1993.

1 female and 1 male colony on a shell of Penion sp. inhabited
by hermit crab, 8.2.1994, Greta Point.

2 female colonies on two gastropod shells inhabited by
hermit crabs, coll. 5.9.1994, Greta Point; colonies removed
from shell and cultivated in small dishes until sporosacs
were released in large numbers (12.9.94).

Description: Hydroid colonies arising from attached,

ramified stolons, erect stems up to 5 mm high, only

occasionally branched once or twice. Stolons and cauli

covered by perisarc which in younger hydranths con-

Fig. 20. Dicoiyne conybearei , from life, a) part of colony with gastrozooids and gonozooid (right); scale bar 1 mm. b) female
gonozooid, CL; scale bar 0.2 mm. c) two free-swimming female sporosacs, upper with one tentaculozooid, lower with two
tentaculozooids; scale bar 50 mm. d) nematocysts: microbasic euryteles of three sizes (desmoneme not shown); scale bar
10 mm.

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tinues over hydranth body as thin pseudohydrotheca
up to tentacle base, in larger hydranths it terminates
at base of hydranth body as a delicate cup-like
expansion. Hydranths of gastrozooids cylindrical with
conical hypostome. Below hypostome one whorl of
12 (11-14) thin filiform tentacles of very different
length, pointing alternately up- and downwards.
Gonozooids arise from stolons or as side branches
from cauli, basal portions covered by perisarc as in
gastrozooids. Body cylindrical with a large distal
swelling studded with nematocysts. No tentacles or
mouth present. Below terminal swelling a zone where
gonophores are produced. Up to 20 or more gono-
phores per gonozooid, but number varies consider¬
ably. Gonophores spherical to oblong. Gonophores
develop into sporosacs which are liberated. Female
and male sporosacs originate on separate colonies.
Liberated sporosacs oblong, up to 0.15 mm, covered
by a ciliated layer of cells which also contains some
nematocysts. This layer surrounds the gametes.
Mature female sporosacs contain one egg only. A
spadix of variable size may be present. In the
population from Wellington the majority of sporosacs
have 2 ciliated tentaculozooids filled with a chordoid
gastrodermis. Some of the sporosacs with one ten-
taculozooid only, or the second one reduced to a
variable degree. Nematocysts:

a) larger microbasic euryteles, in gonozooids, (18-
23) x (6-8) pm, s = 0.8.

b) medium sized microbasic euryteles, in gastro- and
gonozooids, (7-9.5) x (4-6) pm, s = 0.8.

c) smaller microbasic euryteles, on gonophores, (4.5-
6) x (2.5-3.5) pm.

d) desmonemes, in tentacles only, discharged with 4
coils, (4-4.5) x (2-3) pm.

Measurements: gastrozooid hydranths around 1 mm
long and 0.2-0.25 mm diameter; tentacles 1 mm long
and 40-50 mm diameter, gonozooids up to 1.2 mm.

Type Lcxtality: Ireland.

Remarks: Dicoryne conybearei normally liberates sporo¬
sacs having one egg and one tentaculoid only. This
makes them quite distinct from Dicoryne comferta which
has sporosacs with two eggs and two tentaculoids.
The sporosacs of the investigated population from
Evans Bay, however, had one egg and 1-2 tentaculoids
(even within the same colony). In some cases it was
rather difficult to decide whether they were really
2 tentacles or only one together with the elongated
remainder of the original stalk. Ashworth and Ritchie
(1915) also observed rare aberrant sporosacs of D.
conybearei with 2 tentaculoids. Although it may be that
the present population is not D. conybearei, a separation
based on a single, variable character does not seem

justifiable. Thus the population from Wellington was
assigned to D. conybearei. The other characters agree
rather well with the good descriptions given by
Ashworth and Ritchie (1915).

Lendenfeld (1884) described Dicoryne annulata from
the south coast of Australia. Unfortunately, Lenden-
feld's description is entirely unsatisfactory and this
species is at present not recognisable. Therefore it
cannot be compared here to the population from New
Zealand.

Records from New Zealand: Greta Point, Wellington
Harbour (first record by this study).

Other Records: Great Britain; Ireland; Western Medi¬
terranean (Ashford & Ritchie 1915); Japan (Hirohito
1988).

Gravelya Totton, 1930

Colonial hydroids arising from ramified stolons.
Hydrocauli rarely branching, covered by perisarc.
Hydranths fusiform with a dome-shaped hypostome
and with one whorl of filiform tentacles. Hydranth
covered by pseudohydrotheca that does not envelop
tentacles. Gonophores develop on stolons and cauli
and remain fixed as sporosacs.

Type Species: Gravelya antarcticum (Hickson & Gravely,
1907).

Remarks: Generic separation in the families Bougain-
villiidae and Pandeidae is most readily based on their
distinctive medusae. Where this medusa phase is
reduced, systematic difficulties arise because inde¬
pendent reductions within different genera may often
produce almost identical species (an intermediate step
is described by Calder 1993). To fit these groups into
a workable system based on phylogeny seems almost
impossible.

Rees (1938) re-erected the genus Rhizorhagium M.
Sars, 1874 and included in it all unbranched, colonial
polyps with one whorl of filiform tentacles and with
fixed sporosacs. The type species is R . roseum M. Sars,
1874. Similar hydroid genera like Parawrightia Warren,
1907, Gravelya Totton, 1930, and Aselomaris Berrill, 1948
were later also referred to Rhizorhagium (e.g., Millard
1975; Bouillon 1980). Calder (1988) again separated
from this group members with a nipple-shaped
hypostome, namely Parawrightia and the type species
of Rhizorhagium (R. roseum). Both genera were united
by Calder (1988) in a new subfamily Rhizorhagiinae.
Because the reduction of medusae to fixed sporosacs
alone is not a reliable synapomorphy as it may have

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occurred several times independently (see Petersen
1990), such a separation makes sense. The nipple¬
shaped hypostome may indicate an origin other than
from typical bougainvillids. But not all members of
Rhizorhagium sensu Rees (1938) have a nipple-shaped
hypostome and these species must be placed into other
genera. These remaining species are here distributed
into the genera Gravely a Totton, 1930 and Aselomaris
Berrill, 1948. At present, Gravely a is distinguished from
Aselomaris by the development of sporosacs only from
the cauli in the latter species (Berrill 1948). Whether
Aselomaris also has nematocysts on its eggs is not
known. The small difference in the origin of the gono-
phores in these two species makes generic separation
very debatable. At present it seems preferrable to
retain these genera until a broader revision dis¬
entangles the family Bougainvilliidae and allows a
more natural grouping. For this also the genus Garveia
must be considered (this genus is similar to Gravelya,
but has branching stems).

Only Gravelya antarcticum is known from New
Zealand.

Gravelya antarcticum (Hickson & Gravely, 1907)

(Fig. 21a-d)

Perigonitmis antarcticum Hickson & Gravely, 1907: 4, pi. 1,
figs 1-3, pi. 4, fig. 32.

Atractylis antarctica : ?Vanhoeffen 1910: 283, fig. 8.

Gravelya antarcticum : Totton 1930: 139, fig. la-b.
Rhizorhagium antarcticum: Millard 1971: 401, fig. 3A-D.
Perigonimus antarcticus: Stepanjants 1979: 10, pi. 1, fig. 1.

Material Examined:

2 living colonies from Steeple Rock Beacon (Wellington Har¬
bour entrance), 10 m, 30.3.1994. One fertile female colony
covering several square cm on ascidian (Pyura sp.). One
other colony small, on oyster shell, few gonophores
present. Cultivated for few days only.

Description: Hydroid colonies reaching a height of

3 mm, arising from attached, ramified stolons. Hydro-
cauli only rarely branching, covered by a single¬
layered perisarc which extendsas filmy pseudohydro¬
theca over body of hydranth, but does not envelop
tentacles. Perisarc and pseudohydrotheca infested
with detritus.

Fig. 21. Gravelya antarcticum, all from life and CL. a) part of colony with gonophores arising from cauli, hydranth at left
has pseudocapitate tentacles; scale bar 0.2 mm. b) gonophore arising from stolon, same scale as a), c) single hydranth, same
scale as a), d) nematocysts: microbasic eurytele, desmoneme; scale bar 10 Jim.

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Younger hydranths can almost completely retract into
pseudohydrotheca. Hydranths fusiform, up to 0.5 mm
long, with conical hypostome, with one whorl of 6-
12 filiform tentacles. Older hydranths bearing gono-
phores on their cauli often have reduced tentacle
numbers or even reduced hydranths (reproductive
exhaustion). Gonophores arise from stolons and from
cauli, up to 2 gonophores per caulus. Gonophores
remain fixed as sporosacs. Female gonophores oblong
(0,5 x 0.3 mm), covered by thin transparent perisarc,
with long, red spadix, without radial canals or circular
canals. Up to 20 eggs per sporosac, very opaque,
mature eggs covered with a layer of nematocysts.
Nematocysts:

a) microbasic euryteles, (6-8) x (2.5-4) pm.

b) desmonemes, (4-5) x (2.5-4) pm.

Measurements: stolon diameter 60-90 pm; caulus dia¬
meter 70-80 pm.

Type Locality: McMurdo Bay, Antarctica.

Remarks: The available samples of Gravelya antarcticum
were of somewhat shorter stature than the ones des¬
cribed by Hickson and Gravely (1907) and Millard
(1971), but otherwise fit these descriptions well. The
shape of the gonophore seems to be somewhat vari¬
able. A similar species is Aselomnris sagamiense (Hiro-
hito, 1988) n. comb. This Japanese species is char¬
acterised by a double-layered perisarc, which was not
found in the New Zealand material. Hirohito's species
is here assigned to Aselomnris because it develops
gonophores from cauli only.

The covering of the eggs with a layer of nemato-
cytes underlines the close affinity of Gravelya
antarcticum with the genus Bougainvillia , where such a
covering is very frequent. Some of the younger
hydranths showed an unusual behaviour — they were
able to contract the terminal region of the tentacles to
a swelling, thus giving the impression of capitate
tentacles. This pseudo-capitate state was maintained
for quite a long time. Pennycuik (1959) described
similar behaviour in a doubtfully identified Aselomnris
arenosa.

Records from New Zealand: Wellington Harbour (new
record from this study).

Other Records: Antarctica; Marion Island (Prince
Edward Island, South Africa).

Koellikerina Kramp, 1939

Colonial hydroids, erect and branching, covered by
perisarc infested with foreign material. Perisarc also

envelops hydranth body and basal part of tentacles as
pseudohydrotheca. Hydranths with one irregular
whorl of filiform tentacles. Gonophores with a stalk
arise singly from hydrocauli and are liberated as free
medusae. Medusa with eight groups of marginal
tentacles of similar structure. With or without ocelli.
Four dichotomously branching oral tentacles present.
Gonads in perradial or adradial position (Bouillon
1985a).

Type Species: Koellikerina fasciculata (Peron & Lesueur,
1809).

Remarks: The only life cycle known in this genus is
that of the type species (Petersen & Vannucci 1960).
Only one species is known from New Zealand.

Koellikerina maasi (Browne, 1910) (Fig. 22)

Koellikeria maasi Browne, 1910: 22, pi. 4, figs 1-5.

Koellikeria maasi : Vanhoeffen 1912: 361, pi. 25, fig. 2.
Koellikerina maasi : Kramp 1959: 112, fig. 100; Kramp 1961:
85; Kramp 1968: 36, fig. 93; O'Sullivan 1982: 36, fig. 15,
map 14; Bouillon 1995: 229.

Fig. 22. Koellikerina maasi from Stn N465; the length of
the tentacles may not be accurate; scale bar 1 mm.

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c c]Q

Material Examined:

2 medusae from NZOI Stn N465, 47‘40.7'S, 167°01.2'E, 0-
154 m, 5.2.1975, 9 mm diameter.

Description: Medusa with cylindrical bell, top rather
flat, jelly thick, apical jelly twice as thick as lateral,
bell-margin in 8 slight lobes due to furrows at the site
of tentacles. No gastric peduncle present. Manubrium
very large, length half of subumbrellar height, cross¬
shaped in section. With 4 perradial oral tentacles
branching dichotomously 5-6 times and ending in only
very small terminal swellings. Four voluminous
gonads, interraciial in position, covering almost the
whole manubrium, separated perradially, with
irregular vertical folds. Four broad radial canals anci
a thinner circular canal present. Marginal tentacles
arranged in 8 groups, 4 perraciial and 4 interraciial. In
each perradial group 7-9 tentacles, in the interraciial
groups ca. 6. Tentacles without bulbs anci of varying
length. The tentacle in the middle of each perraciial
group is longer anci about twice as thick as the others.
Polyp stage unknown.

Type Locality: McMurdo Sound, Antarctica.

Additional Information: There are no ocelli present anci
the stomach has a reci colour in life (Vanhoeffen 1912).

Remarks: The gonads have previously been described
as smooth, but in the present samples they had
irregular vertical folds. Some of these may have been
caused by the fixation however.

Records from New Zealand: Between Stewart Island
anci Snares Islands (Bouillon 1995).

Other Records: Antarctica.

Family HYDRACTINIIDAE L. Agassiz, 1862

Hydroid colonies with sessile hydranths arising either
from a reticulate hydrorhiza with stolons covered
by perisarc, or from a hydrorhiza consisting of an
encrusting layer of coenosarc not covered by perisarc,
or from a calcified encrusting hydrorhiza. Chitinous
spines, tubes, or calcareous spines may be present.
Hydranths polymorphic with gastrozooids, dactylo-
zooids anci gonophore bearing gonozooicis. Gastro-
zooicis with one to several filiform tentacles below
hypostome. Dactylozooicis may be present, either
without or with capitate tentacles. Reproduction by
sessile sporosacs or free meciusae. Exceptionally,
sporosacs may be prociuceci by the hydrorhiza or in
specialiseci cavities of the calcareous skeleton.

Meciusae with or without a gastric peciuncule, mouth
with four simple or ramifieci lips with terminal
nematocyst clusters. With four raciial canals anci
circular canal. The gonacis ciifferentiate on the manu¬
brium in interraciial positions, occasionally they
extenci onto the proximal part of the raciial canals.
Exceptionally, the gonacis can be in an aciraciial
position (genus Hansiella). Marginal tentacles solid anci
not in groups. Ocelli present or not.

Remarks: Bouillon's (1985a) diagnosis is here amended
to accommociate Fiordlandia n.gen. The emenciation
concerns the skeletal structures: not only spines are
possible, but also tubes like the protective stolons seen
in Fiordlcindici. Fiordlcindici is placeci in the Hyciractini-
iciae on account of its sessile, ciimorphic zooicis.

It is rather obvious that the family Hyciractiniiciae
is in neeci of revision. Generic limits should be re¬
defined baseci on phylogeny, which may be rather
complicated anci ciifficult to assess with morphological
methocis (cf. Cunningham & Buss 1993). However, the
present publication cannot deal with such a major
revision and a provisional system as ciefined by
Bouillon (1985a) is useci.

Characteristics of New Zealand genera:

Hy dr actinia: sessile sporosacs, with encrusting
hycirorhiza not covereci by perisarc.

Podocoryna: with a free, well-developed meciusa
having oral lips stucicieci with nematocysts.
Stylactaria: sessile sporosacs or ciegenerate medusae,
anci reticulate hycirorhiza covereci with perisarc.
Fiordlandia : sessile sporosacs, gastrozooicis with
tentacles scattereci in region below hypostome, anci
erect tubular stolon-like structures.

Hydractinia van Beneden, 1841

Colonial hydroids with hycirorhiza composeci initially
of tubes covereci with perisarc which form a tight
meshwork anci ultimately coalesce into an encrusting
layer covereci by nakeci coenosarc (basal plate). This
layer often forms spines which can be simple or
brancheci. Hydranths are polymorphic with gastro¬
zooicis, gonozooicis, anci ciactylozooicis. Gastrozooicis
with one or two closely set whorls of filiform tentacles.
Dactylozooicis of various types may be present, lacking
mouth anci tentacles. The gonophores remain fixeci as
sporosacs anci arise from normal hydranths, reciuceci
hydranths, or ciirectly from the hycirorhiza (Bouillon
1985a).

Type Species: Hydractinia lactea van Beneden, 1844.
Remarks: Hydractinia is often rather ciifficult to ciis-

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tinguish from Styl act aria (p. 54) because the encrusting
layer without a perisarc covering is only formed in
well-grown colonies. Therefore, only fertile material
can be readily identified.

Many Hydractina species live on gastropod shells,
especially ones inhabited by hermit crabs. This associ¬
ation may be a symbiosis as shown by Brooks and
Gwaltney (1993). Hydractinia species can form sibling
species which may not be identifiable using ordinary
morphological characters (Buss & Yund 1989). As
some of these cryptic species may be host-specific for
one hermit-crab species, the crab species hosting the
Hydractinia colony should be identified if possible. An
excellent key to the hermit crabs of the Otago region
is provided by Schembri and McLay (1983). Their key
is also useful for the remaining parts of New Zealand.

Ralph (1961c) recorded a Hydractinia species from
the Chatham Islands. Ralph's material was infertile
and therefore not identifiable. This record is not treated
here again.

Characteristics of species present in New Zealand:
Hydractinia parvispina: on rocks, with around 8 eggs
in female gonophores.

Hydractinia novaezelandiae: on hermit-crab shells or
other substrata; with branching spines, 2 eggs in
female gonophores, and small hydranths.
Hydractinia rubricata: on shells inhabited by hermit
crabs; with groups of spines on elevation of basal
plate, basitrichous anisorhiza, with 5-7 eggs in
female gonophore, and large hydranths.

Hydractinia parvispina Hartlaub, 1905 (Fig. 23a-b)

Hydractinia parvispina Hartlaub 1905: 518, fig. a; Jaederholm
1905: 5, pi. 3, figs 4-5; Ralph 1953: 63, fig. 1; Millard 1971:
402, fig. 4; Stepanjants 1979: 14, pi. 1, fig. 8.

Material Examined:

No material seen.

Description (After Millard 1971): Hydroids colonial,
growing mostly on rocks or on gastropod shells.
Hydrorhiza covered with a layer of naked coenosarc.
Spines smooth and hollow, closed or open at the tip,
the largest reaching 0.5 mm in height but most of them
smaller. Polyps polymorphic with gastrozooids and
gonozooids; no tentaculozooids or spiral zooids
present. Gastrozooids in life up to 6 mm high, with
11-16 tentacles, which in contracted individuals
appear to arise in 2 alternating whorls. Gonozooids
smaller than gastrozooids, up to about two-thirds of
the size of gastrozooids, with well-developed conical
hypostome and 4-6 tentacles. Male gonophores borne
in a circle on distal half of gonozooid, usually about 8

Fig. 23. Hydractinia parvispina after Ralph (1953), redrawn
with kind permission of the publisher, a) colony with female
gonozooid, tentaculozooid and spine; scale bar 0.5 mm. b)
male gonophore; scale bar 0.2 mm.

large gonophores per gonozooid, but up to 18 have
been found including immature buds. Male gono¬
phores subspherical, with short pedicel and no radial
canals, reaching a maximum size of 0.41 x 0.37 mm.
Nematocysts:

a) elongated capsule, 9.5 x 3.6 pm.

b) small bean-shaped capsules, 6x3 pm.

According to Hartlaub's (1905) description, the

female gonophores have a number of eggs. Judging
from his figures (Fig. A, C) they must contain at least
6, probably more. According to Ralph (1953) they have
approximately 8 eggs.

Type Locality: Tierra del Fuego and the Falkland
Islands.

Remarks: Hydractinia parvispina is not a well-defined
species. The best description was given by Millard
(1971), but details on nematocysts and growth sub¬
strates are inadequately known. Ralph (1953) reported
this species with a query from Portobello where it was
growing on rocks. Although Millard (1971) considered
this record as doubtful, Ralph's (1953) short diagnosis
and figure are compatible with the present concept of
H. parvispina as given by Hartlaub (1905) and Millard
(1971). The major discrepancy is the presence of ten¬
taculozooids in the material from New Zealand.
However, the presence of tentaculozooids is not
diagnostic for individual colonies as in other species
they may also be absent in a majority of colonies. The
best support for Ralph's material having been H . parvi-

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@®®C

spina is its occurrence on intertidal rocks. The sampling
site of Ralph's material (intertidal near Portobello
Marine Biological Station) was re-examined for this
study, but no Hydractinia species could be found

at that time. The only abundant hydractiniid found
was Stylactaria otagoensis n. sp. (see p. 54). Hydractinia
parvispina and Stylactaria otagoensis appear very
similar, differing mainly in the structure of the hydro-

Fig. 24. Hydractinia novaezelandiae n. sp., from preserved type material, a) part of colony with gastrozooids, gonozooids and
spines; scale bar 0.2 mm. b) two typical spines showing distal branching, scale bar 0.2 mm. c) female gonozooid, same scale as
b). d) female gonophore, spadix shaded; scale bar 0.1 mm. e) male gonophore, with distal patch of nematocysts, same scale as
d). f) nematocysts: microbasic eurytele from hypostome, heteroneme from tentacles, desmoneme; scale bar 10 Jim.

)®®m

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rhiza only. The naked coenosarc in Hydractinia is not
always easy to see and it may be that Ralph actually
observed S. otagoensis. The presence of tentaculozooids
argue in favour of this interpretation. At present, it
seems better to maintain Ralph's identification until
more observations can either confirm or exclude the
presence of Hydrnctinin parvispina in New Zealand.

Records from New Zealand: Intertidal reef near
Portobello Marine Biological Station.

Other Records: Tierra del Fuego, Falkland Islands,
South Georgia, Paulet Island (Graham Land), Prince
Edward Islands.

Hydractinia novaezelandiae n.sp. (Fig. 24a-f)

Material Examined:

Holotype from NZOI Stn Q725 (off Westport), 4.3.1982,
dredged material from 35 m, colonies (male and females)
on shell of Amnldn australis, ~2 cm shell size; colony also
in siphonal canal therefore most probably not inhabited
by gastropod at time of sampling, but no hermit crab
visible either. Deposited as holotype H-646.

NZOI Stn Q726, 41°25.50'S, 172°01.20 , E, dredged from 47 m,
5.3.1982, on ~5 mm fragment of calcareous tube, colony
infertile. Deposited as paratype P-1072.

NZOI Stn J899, 35°510'S, 174°28.0’E (Bay of Islands),
collected intertidally, 22.1.1976, fertile male colony on shell
of Amalda australis, shell empty. Deposited as paratype P~
1079.

Description: Hydroid colonies with hyarorhiza
forming a network of perisarc-covered tubes which is
open at the colony periphery but coalesces towards
the centre to form an encrustation covered with a layer
of naked coenosarc, bearing spines, gastrozooids,
gonozooids and tentaculozooids. There are regularly
scattered knobs in the basal plate. Spines elevated up
to 0.5 mm, distal parts smooth, basal parts with smaller
spines and prickles. The spines are either simple, but
often distally branched with up to 8 ends. Coenosarc
may cover a large part of the spines but polyps do not
arise from them. The morphology and size of spines
depend on their location on the shell. Gastrozooids
reaching up to 1 mm in height (preserved material),
with 4-8 tentacles in one whorl. Gonozooids shorter
than gastrozooids, about as high as largest spines, with
3-4 tentacles, these short and stubby; with up to 8
gonophores in one whorl. Gonophores without radial
or circular canals. Nematocysts present in a distal
patch in gonophores of both both sexes. Female
gonophores with 2 eggs and asymmetrically placed
spadix. Males without spadix. Tentaculozooids
present at shell lips, 0.4 mm long, with distal swelling.

Nematocysts:

a) microbasic euryteles (?) from hypostome and body,
(7-8) x (3-3.5) pm.

b) heteronemes from tentacles, (6-7) x (2.5-3) pm.

c) desmonemes, (3.5-4) x (2-2.5) pm.

Colours: spines amber, otherwise no original colours
preserved.

Type Locality: 42°25.5’S, 171°05.5’E (off Westport),
depth 35 m.

Etymology: The species name refers to New Zealand
where this species seems to have a quite broad dis¬
tribution.

Remarks: The distally branching spines of H. novae¬
zelandiae make this species rather distinct. Similar
spines are known from H. echinata (cf. Broch 1916).
But H. echinata has gonozooids without tentacles. The
spines are also somewhat similar to those of H. mbri-
cata , but in the latter species the spines are more like
tufts on a elevation of the basal plate, while in H.
novaezelandiae they resemble more large spines with
several endings. Besides this difference, H. novae¬
zelandiae and H. rubricata are distinct in many other
characters: overall size of polyps, tentacle numbers,
number of eggs in female gonophores, spadix of male
gonophores. Also the nematocysts are different, but
because only preserved samples were available for H.
novaezelandiae, the nematocysts could not be examined
precisely.

Records from New Zealand: Off Westport and Bay of
Islands.

Hydractinia rubricata n. sp. (Fig. 25a-c)

Material Examined:

3 living colonies on shells inhabited by Pagurus rubricatus
(Hendersen, 1888), collected by dredging, 100 m, off Kai-
koura Peninsula, 4.5.1994. One male colony on a - 5 cm
large shell of Austrofusus glans (Gastropoda) was selected
as holotype and deposited as H-647. The other two, both
females, one on shell of A glans, the other on Argobuccinum
tumida (Gastropoda) were deposited as paratypes P-1074
and P-1075.

3 living colonies in public aquarium, Portobello Marine
Laboratory (Dunedin), May 1994, collected near Dunedin,
shell size 6-7 cm, all inhabited by P. rubricatus, all three
fertile males. Several other hermit crab species were
examined, but none had hydroids on them.

Description: Hydroid colonies epizooic on shells
inhabited by hermit crabs. Hydrorhiza forms network
of perisarc-covered tubes which is open at the colony
penpAeiy but coalesces towards the centre to form an

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encrustation covered with a layer of naked coenosarc,
bearing spines, gastrozooids, gonozooids, and spiral
zooids. There are regularly scattered prickles in the
basal plate. Spines elevated up to 1.5 mm, distal parts
smooth, basal parts prickly as basal plate. The spines
are either single or often grouped together. Very often
they are grouped on a conical elevation of the basal
plate which may also bear polyps. The type and size
of spines depends on their location on the shell.

Gastrozooids reaching in life up to 10 mm in height,
slender, with 16-20 tentacles in one whorl. Tentacles
long, thickest part somewhat distal from origin.

Gonozooids much shorter than gastrozooids, about
as high as spines, with up to 4-6 (max. 10) tentacles,

these short and stubby; below tentacles a zone with
1-4 gonophores in one whorl. Gonophores without
radial or circular canals, with darkly coloured spadix.
Nematocysts mostly absent in gonophores. Female
gonophores with 5-7 eggs with obvious germinal
vesicle. Female gonozooids have a belt of very con¬
spicuous oogonia in the gastrodermis of the gonophore
budding region. Spiral zooids are present along outer
lip of shell opening, isodiametric and expandable. They
often uncoil and recoil synchronously. Nematocysts:

a) microbasic euryteles, (7.5-13.5) x (3-5.5) jam, s = 1.

b) basitrichous anisorhizas, on hydranth body, (6-
8) x (2-2.5) urn.

c) desmonemes, discharged with 4 coils (5-8) x (3-
5) jam.

Fig. 25. Hydractinia nibricata n. sp., all from life, a) part of paratype colony on outer lip of shell opening with gastrozooids,
gonozooids, spiral zooids and spines; scale bar 0.5 mm. b) female gonozooid with two gonophores and oogonia visible in
budding zone; scale bar 0.2 mm. c) nematocysts in pairs of native and discharged: microbasic euryteles, basitrichous
anisorhizas, desmonemes; scale bar 10 pm.

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Type Locality: 42°28 , S / 173°42E (off Kaikoura).

Etymology: The species name is derived from the
hermit crab host Pagurus rubricatus.

Remarks: Hydrachma rubricata is quite distinct from all
other better-known species of the genus. The
basitrichous anisorhiza is especially unusual. Unfortu¬
nately, the nematocysts of few species are known (cf.
Bouillon 1985a). Also the conspicuous oocytes in the
gastrodermis of the gonophore budding region are
unusual. Although in hydrozoans the germ cells are
located in the epidermis, they may originate in the
gastrodermis as in the other cnidarians (Schuchert
1993).

All examined living colonies were found on shells
inhabited by the hermit crab Pagunis rubricatus (Hen-
dersen, 1888), which has a wide distribution in New
Zealand, mostly between 20 and 200 m (Schembri &
McLay 1983). Other living hermit crabs examined from
the same catches that yielded P. rubricatus did not bear
hydroids. But due to the few samples examined, an
association with other species cannot be ruled out at
present.

The spine morphology and especially their size
varied considerably, depending on their position on
the shell. The region adjacent to the inner lip of the
shell-operdng had the largest spines. This is also the
region which is presumably most exposed to mechan¬
ical impacts from the crab and the bottom. The spines
may therefore protect the gonozooids from abrasion.
It would be worthwhile to test this hypothesis with
experiments.

The function of the spiral zooids is unclear. They
were quite regularly found along the outer lip of the
shell opening (Fig. 25a). Many of them uncoiled and
rapidly recoiled synchronously. Spiral zooids occur
only in hydractiniids associated with hermit crabs (cf.
Mills 1976a).

Records from New Zealand: Kaikoura and Dunedin.

Podocoryna M. Sars, 1846

Colonial hydroids with hydrorhiza composed of
perisarc-covered tubes forming a tight meshwork
which may persist as such, or coalesce into a mesh-like
encrusting layer which may be covered by naked
coenosarc, or occasionally by a very thin periderm.
The hydrorhiza can form small spines. Hydranths are
polymorphic with gastrozooids, gonozooids, and
occasionally dactylozooids. The gonozooids are mostly
similar to gastrozooids, but generally smaller and often
with reduced number of tentacles. Gonophores arise

below the whorl of tentacles and are generally covered
by a thin film of periderm; gonophores liberated as
free medusae. Medusae with four or more solid,
simple marginal tentacles, never in groups. Marginal
bulbs present, with or without ocelli. There are four
or eight oral arms which are outgrowths of the lip only;
they are simple or branched and provided with a tight
cover of nematocysts. Four radial canals and circular
canal always present. The gonads are interradial,
occasionally extending on proximal parts of radial
canals. In some species the medusae may reproduce
asexually by budding new medusae (Bouillon 1985a).

Type Species: Podocoryna came a M. Sars, 1846.

Remarks: The above diagnosis certainly does not reflect
a monophyletic group. The presence of a medusa must
be considered a plesiomorphy that cannot reveal a
monophyletic taxon. The encrusting layer of hydro¬
rhiza might be one of the few convincing synapo-
morphies within the Hydractiniidae, which is,
however, not used by the present system to delimit a
taxon. Thus, grouping together species with and
without such an encrusting layer generates a para-
phyletic taxon. A future revision of the Hydractiniidae
might thus abandon this genus, as proposed, for
example, in Broch (1916) and Naumov (1969) (cf. also
Cunningham & Buss 1993). For the same reasons as
given in the family diagnosis, however, a revision is
not made here. Calder (1988) showed that the correct
spelling is Podocoryna, and not the more frequently
used Podocoryne.

To identify a Podocoryna colony correctly, the adult
medusa stage must be grown from them (except for
P. bella).

Characteristics of New Zealand species:

Podocoryna bella: polyps growing on Pigfish ( Congio-
podus leucopaecilus), hydrorhiza reticulate, adult
medusa unknown.

Podocoryna minima: polyp stage unknown, medusae
with medusae buds on manubrium, with peduncle,
and four marginal tentacles.

Podocoryna minuta : polyp stage unknown, medusae
with medusae buds on manubrium, with peduncle,
and eight marginal tentacles.

Podocoryna australis: polyps with encrusting
hydrorhiza, adult medusa with 8-14 marginal ten¬
tacles, no medusae buds on manubrium, and with¬
out distinct peduncle.

Podocoryna bella Hand, 1961 (Fig. 26a-d)

Podocoryne bella Hand 1961: 91, fig. 1; Kramp 1968: 29, fig.
73.

0®gx

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Material Examined:

Preserved fin of Pigfish ( Congiopodus leucopnecihts) with P.
belln collected and identified by C. Hand near Portobello
Marine Laboratory, 12.1.1960, colony without medusae
buds, material kept by Portobello Marine Laboratory
(•unedin). Type material could not be obtained.
Preserved medusae kept by Portobello Marine Laboratory:
label reads: young medusae of Podocoryne belln from pig-
fish from Aquarium (collected Otago Harbour), released
24-29 October, fixed 2.11,1962. Collector unknown.

Description:

Polyp stage (from own observations and Hand 1961) :
Hydroid colonies growing on Pigfish ( Congiopodus
leiicopnecilus), with gastrozooids and gonozooids.
Hydranths sessile, arising from ramified, loosely
adhering stolons. Stolons covered by very thin (1 pm)
perisarc only, not forming a basal plate, polyps without
perisarc cup at base. Spines or dactylozooids not
observed. Gastrozooids up to 2 mm high, columnar,
with conical hypostome. With one whorl of 10-12 (6-
15 range) filiform tentacles. Gonozooids scattered in
centre of colony, similar to gastrozooids but about half
the size and with 6-8 tentacles only. Gonophores (1-
7) in the middle of hydranth body. Gonophores are
released as free medusae with 8 tentacles.
Nematocysts:

a) microbasic euryteles (?), some on body but con¬
centrated on hypostome, not seen discharged, (8-
9.5) x (3-3.5) pm.

b) smaller microbasic euryteles, on tentacles, (6-
7) x (2-3) pm, s = ~0.7.

c) desmonemes, discharged with four coils, (5-6) x (2-
3) Mm.

Colour: pale, translucent white.

Young medusae: Hemispherical, 0.8 mm high and
1 mm broad, with rather thin jelly (0.08 mm) of
uniform thickness. No peduncle present. Manubrium
length half of bell cavity, cylindrical, mouth with 4
perradial clusters of elongated euryteles. Lips not
elongated to oral arms. No incipient gonads visible.
Four radial canals and ring canal present, these rather
thin. Four perradial tentacles and 4 interradial
tentacles, all of similar length. All 8 tentacles with a
tentacle bulb. No ocelli observed.

Adult medusa: Unknown.

Type Locality: Otago Harbour.

Remarks: Podocoryna belln is not a sufficiently described
species. The adult medusa remains unknown and there
are also not enough data available on its ecology. It
would be very important to know whether Congio-
podus leucopaecilus is the sole host or not. Congiopodus
can support quite a number of epizooic animals.
Several Pigfish were examined alive for this study.
Although none had P. belln , some were found to bear
rather large colonies of the thecate hydroid Phinlelln
sp. and also bryozoans. As noted by Hand (1961)
Turritopsis also occurs on Pigfish. Therefore, P. belln
might not be restricted to Congiopodus and may occur
on other substrata too. The medusa resembles that of
Podoconjnn nustrnlis (p. 51). In the available medusae
of P. belln, however, the jelly was thinner and there

Fig. 26. Podocoryna bella from preserved samples, a) gastrozooid and hydrorhiza, CL; scale bar 0.2 mm. b) gonozooid with
medusa buds, redrawn from Hand (1961) with kind permission of the publisher, same scale as a), c) nematocysts of polyp
stage: microbasic euryteles and desmoneme; scale bar 10 pm. d) 6 to 10-day old medusa; scale bar 0.2 mm.

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was no trace of gonads visible. This may also result
from cultivation. More data are needed for a discus¬
sion of the identity of P. bella and its relation to other
Podocoryna species growing on fish (cf. Komai 1932;
Hirohito 1988).

Records from New Zealand: Only known from Otago
Harbour.

Other Records: Not known outside New Zealand.

Podoconjna minima (Trinci, 1903) (Fig. 27)

Cytaeis minima Trinci, 1903: 1, pi. 1, figs 1-30.

Podocoryne simplex Kramp, 1928: 45, fig. 20.

Podocoryne minima : Russell 1953:134, figs 63-64; Kramp 1961:
69; Kramp 1968: 28, figs 69-70; Brinckmann-Voss 1970:
pi. 7, fig. 1; Uchida & Sugiura 1977: 53, figs 12; Barnett
1985: 81, fig. 8D, pi. 3, fig. C; Goy et al 1991: 107, fig. 17.

Material Examined:

No material seen: identified and described after photograph
and figure in Barnett (1985).

Description: Medusa with globular bell, 0.3-0.9 mm
high. Apical jelly slightly thickened. Manubrium on a
well-developed peduncle. Manubrium length half of

Fig. 27. Podocoryna minima , drawn after photograph from
Barnett (1985); scale bar 0.1 mm.

bell cavity, cylindrical with perradial lip margins
elongated to oral arms terminating in a spherical knob
of nematocysts. With interradial medusae buds on
manubrium. Four radial canals and ring canal, rather
narrow. With 4 perradial tentacle bulbs and tentacles.
No ocelli present.

Polyp stage unknown.

Type Locality: Gulf of Naples, Mediterranean.

Remarks: No material of this species could be
examined, but the photograph and figure given by
Barnett (1985) are clear enough to recognise this
species. A detailed description including also develop¬
mental stages are given by Uchida and Sugiura (1977).

From the literature, gonads are initially interradial,
but when mature encircle the manubrium. The size of
sexually mature medusae is not different from asex-
ually reproducing ones. The colour of marginal ten¬
tacle bulbs and stomach yellowish-brown or brown
(Russell 1953; Uchida & Sugiura 1977).

The descriptions of Podocoryna minima (Trinci, 1903)
and Podocoryne simplex Kramp, 1928 are very similar
and both species are not objectively separable at
present. Therefore, following Uchida & Sugiura (1977)
P. simplex is here also included in the synonymy of P.
minima . Podocoryna minima is a very simple medusa
without many distinctive characters. However, it may
well be that characters of the polyp stage will allow a
separation of the two species as soon as their life cycles
become known.

Records from New Zealand: Leigh Marine Reserve
(Barnett 1985).

Other Records: Mediterranean, Great Britain, Japan
east coast of USA.

Podocoryna minuta (Mayer, 1900) (Fig. 28a-b)

Dysmorphosa minuta Mayer, 1900b: 41, pi. 18, fig. 42.
Podocoryne minuta: Mayer 1910: 140, pi. 14, fig. 1; Kramp
1959: 102, fig. 68; Kramp 1961: 69; Goy 1972: 978; Goy et
al 1991: 107, fig. 18.

Material Examined:

4 medusae collected by T. Barnett in Leigh Marine Reserve
1984, all animals without mature gonads but all budding
medusae, one animal deposited.

Description: Medusa with bell-shaped umbrella, 0.5-
0.7 mm high, slightly higher or as high as broad. With
apical projection. Manubrium on well-formed
peduncle. Manubrium length half of subumbrellar
height, simple, cylindrical. Four perradial lips elon-

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Fig. 28. Podoconjna minuta after preserved specimens from
Leigh, a) Medusa with medusae buds; scale bar 0.1 mm. b)
nematocysts: microbasic eurytele, desmoneme; scale bar
10 mm.

gated to tentacle-like oral arms terminating in spherical
knobs with nematocysts. Medusae buds on inter-
radial wall of manubrium. With narrow radial canals
and ring canal. Four perradial and 4 interradial ten
tacular bulbs and tentacles present. All 8 tentacles
flexible and of equal length, evenly covered with
nematocysts. No ocelli present. Newly budded
medusae are released with 8 tentacles. Nematocysts:

a) microbasic euryteles, on tentacles and mouth
clusters, (5.5-6.5) x (2.5-3) pm, s = 0.9.

b) desmonemes, (3.5-4) x (1.5-2.5) pm.

Polyp stage unknown.

Type Locality: Tortugas, Florida.

Remarks: According to Goy (1972) the gonads may
mature even with continued medusae budding. The

gonad encircles the manubrium below the medusae
buds. Podoconjna minuta from New Zealand is larger
than described by Mayer (1900b) (up to 0.7 versus
0.3 mm). But as it otherwise agrees well with other
descriptions, and size alone cannot be used for a
specific separation, the population from New Zealand
is here assigned to this species. Goy et ai (1991) even
reported a maximal size of 2 mm for this species.
Another similar species is P. tenuis (Browne, 1902) from
the Falkland Islands, but this species has very short
oral arms. More information on life cycles and ecology
is needed for all populations for properly delimiting
the species.

Podoconjna minuta is a rarely reported species with
a patchy distribution, but it may often have been over¬
looked owing to its small size.

Records from New Zealand: Leigh Marine Reserve
(Barnett 1985, as Podoconjne sp. 1).

Other Records: Florida, West Africa, Mediterranean
(Kramp 1961).

Podoconjna australis n.sp. (Fig. 29a-g)

Material Examined:

1 colony on shell of hermit crab, collected 16.12.1993, Evans
Bay, depth 1 m, with dactylozooids, medusae grown to
maturity (male), material deposited.

10 colonies on shells of living Cominella maculosa (Gastro¬
poda), collected 31.12.1993, Narrow Neck Beach, Auck¬
land, 1 m, medusae grown to maturity. One female
medusa with 13 tentacles deposited as holotype H-649,
remaining medusa deposited as paratype P-1077, 4 shells
with polyp colonies deposited as paratype P 1078.

3 colonies on shells, one with gastropod, two with hermit
crabs, collected 28.1.1994, 0.3-1 m, Mahanga Bay, Welling¬
ton Harbour, one infertile.

2 colonies on shells with hermit crabs, collected 8.2.1994,
Evans Bay, depth 0.5 m, fertile.

Approx. 12 adult medusae from plankton sampled in Evans
Bay from November 1993 to January 1994, up to 16
tentacles, medusae spawned and produced planulae,
some material deposited as paratype P-1079.

Several medusae collected by T. Barnett from Leigh Marine
Reserve, 1983-1984, labelled Podocotyne sp. 2.

Several living polyp colonies and medusae of P. carnea
originating from the Mediterranean and Atlantic, for com¬
parison.

Description:

Polyp stage: Hydroid colonies growing on gastropod
shells inhabited by gastropods or hermit crabs.
Hyciranths arise from encrusting hycirorhiza covered
with a layer of naked perisarc, hydranths with no basal
collar of perisarc. Hydranths polymorphic with larger
gastrozooids, smaller and thinner gonozooids, and

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rarely with tentaculozooids. Gastrozooids tubular to
fusiform, shape varies considerably in samples from
nature, 1-4 mm high (normally 2 mm; 0.3-0.4 mm
diameter), with bluntly conical hypostome, 10-14
(max. 20) filiform tentacles of unequal length in one
whorl below hypostome. Gonozooids 0.8-2 mm high,
with 3-6 tentacles and somewhat below to them a zone
of medusae buds (1-8 buds). Tentaculozooids rare,
only seen in specimens from hermit crabs, long and
slender, very extensible, thickened at origin. Spines
present, 0.2-0.6 mm high, with smooth surface, amber
coloured, not in groups. Nematocysts:

a) larger microbasic euryteles, mainly on body, (11-
12) x (3-5) pm, s ~1.

b) smaller microbasic euryteles, mainly in tentacles,
(8.5-9) x (3-4) pm.

c) desmonemes, in tentacles, four coils when dis¬
charged, (5.5-7) x (3-4) pm.

Newly liberated medusa: Spherical, 0.6-0.8 mm high,
with thin mesogloea, apical canal, no peduncle. Manu¬
brium tubular, length approximately half of bell height,
with 4 perradial nematocyst clusters on mouth margin.
Incipient gonads sometimes visible in females (at this
stage egg size around 40 pm). Four radial canals ending
in large marginal bulbs and 4 perradial tentacles and
0-4 shorter interradial tentacles with bulbs. Tentacles
directed upwards when contracted. Mostly no or only
very few (< 5) scattered nematocysts on exumbrella.
No medusa budding during later development.
Nematocysts:

a) microbasic euryteles of tentacles, (7-8.5) x (3-

3.5) pm, s ~1.

b) microbasic euryteles of mouth clusters, (9.5-
11) x (3-3.5) pm, s ~1.

c) desmonemes from tentacles, discharged with four
coils, (5-6.5) x (3-4) pm.

Adult medusa: With bell-shaped umbrella, up to
1.6 mm, higher than wide to as wide as high, jelly
thicker at apex, velum when dilated half radius width,
slight peduncle (up to one-tenth of manubrium length)
may be present or not. Manubrium tubular, length half
to two-thirds of bell cavity, with 4 simple perradial
clusters of elongated euryteles at mouth margin.
Gonads interradial. Four radial canals ending in
conspicuous bulbs. Four interradial bulbs mostly

present. Tentacle number normally 10-14 (8-16 range).
Nematocysts:

a) microbasic euryteles of tentacles, (7-7.5) x (3-

3.5) pm.

b) microbasic euryteles of lip clusters, (9-10) x (2.5-

3.5) pm.

c) desmonemes, (4.5-5) x (2.5-3) pm.

Size of spawned eggs (from medusa grown from
polyps): 131 pm (s.d. 6.8 pm, n = 7).

Type Locality: North end of Narrow Neck Beach,
Devonport, Auckland.

Etymology: The species name australis, Latin, southern,
was chosen to contrast the species from similar ones
occurring in the northern hemisphere.

Remarks on Life Cycle: Medusae released from colonies
collected at Wellington and Auckland were grown to
maturity (spawning), attained after one week. The
shape of the bell was somewhat variable, but corre¬
sponded to medusae found in the plankton. Tentacle
numbers in mature cultivated specimens were 8-13
(Fig. 29c). Fertilised eggs from medusae found in the
plankton resulted in many planulae which, however,
failed to metamorphose.

Remarks on Ecology: Podoconjna australis polyps were
found in shallow water (1-2 m) on shells of Cominella
maculosa (Gastropoda) living on a sand/mud bottom
(Auckland, Wellington), and also on other shells
inhabited by hermit crabs (Wellington Harbour).

Remarks: Podoconjna australis closely resembles P.
came a M. Sars, 1846, P. selena Mills, 1976b, and P.
hyamensis Hirohito, 1988. The only character distin¬
guishing the latter from P. australis and the other
species is the granular surface of its spines. Podoconjna
cornea as defined by Edwards (1972) is very variable,
its medusa forming a morphological cline from
northern Europe to the Mediterranean, varying in size,
tentacle number, and timing of maturity (cf. Benard-
Boirard 1962). The polyp stage shows considerable
variation in size and proportions depending on growth
condition (unpublished observations on cultures).
Podoconjna cornea, however, very rarely has more than
8 tentacles and never more than 10 (Edwards 1972).

Fig. 29. (opposite) Podoconjna australis n. sp., all from life, a) part of a polyp colony with spines, gastrozooids, gonozooids
with medusae buds, and tentaculozooids (right margin). Gastrozooids are shown in various degrees of contraction and
stomach filling; scale bar 1 mm. b) newly released medusa with 6 tentacles; scale bar 0.5 mm. c) cultivated 8-day old female
medusa which was able to spawn, same scale as b). d) adult female medusa from plankton, not all tentacles shown, the
slight peduncle shown in this specimen is not always present; same scale as b). e) manubrium of an adult male medusa from
the plankton, same scale as b). f) nematocysts of polyp: two sizes of microbasic euryteles and desmonemes; scale bar 10 jam.
g) nematocysts of medusa: microbasic euryteles, desmonemes, microbasic eurytele of lips; scale bar 10 (am.

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In addition, its exumbrella at liberation is covered with
dozens of nematocysts, and a peduncle may be present
only during early development Podoconjna australis
in contrast has regularly more than 8 tentacles and
often more than 10; at liberation there are no or < 5
nematocysts on the exumbrella and occasionally a
shallow peduncle maybe present in mature animals.
Podocoryna selena from Florida is even more similar to
P. australis because the former has 8-14 tentacles and
lacks nematocysts on the exumbrella at liberation. The
only difference between the two species is that in P.
australis the gonads are not developed at liberation
and it may reach a 16-tentacle stage. The male gonads
of P. australis are even invisible just after liberation
and develop only later.

Podoconjna carnea, P. selena , and P australis polyps
are not distinguishable from existing descriptions.
Admittedly, the differences between the medusae are
small and splitting into different species is debatable.
The short life span of Podoconjna medusae (ca. three
weeks) and the wide geographic separation (North
Atlantic, Florida, New Zealand) make it more plausible
that these populations do not belong to the same
species. Some initial molecular analysis (Cunningham
& Buss 1993) indicate that P selena and P carnea show
a few differences. It would be very rewarding to
examine the systematics of hydractiniids by methods
used for population genetics (e.g., Thorpe et al. 1992;
Thorpe & Sole-Cava 1994).

Records from New Zealand: Polyp from Auckland and
Wellington Harbours. Medusa from Goat Island, Leigh
(Barnett 1985, as Podocoryne sp. 2), and Wellington
Harbour (this study).

Stylactaria Stechow, 1921

Colonial hydroids with hydrorhiza formed by stolons
that are completely covered by perisarc, no layer of
naked coenosarc, with or without spines. Polyps poly¬
morphic with gastrozooids, gonozooids, and some¬
times dactylozooids. Gastrozooids with one or rarely
several whorls of filiform tentacles. Gonozooids simi¬
lar to gastrozooids but often smaller and with reduced
number of tentacles; gonophores either remain fixed
as sporosacs or are liberated as degenerated medusae.
Medusae have four simple radial canals and an
elongated manubrium without mouth opening; no
oral arms with nematocysts; gonads surround the
manubrium. With four to eight marginal bulbs,
exceptionally with marginal tentacles (after Bouillon
1985a).

Type Species: Stylactaria inermis (Allman, 1872).

Remarks: Ihe genus Stylactaria has been reviewed by
several authors (Iwasa 1934; Bouillon 1971; Namikawa
1991). The generic name was recently modified by
Calder (1988) from the commonly used Stylactis, which
was shown to be a synonym of Hy dr actinia, to Sty¬
lactaria.

There is only one species of Stylactaria known from
New Zealand.

Stylactaria otagoensis n. sp, (Fig. 30a-d)

Material Examined:

Holotype H-651 (NIWA), part of larger female colony grow¬
ing on holdfast of Macrocystis pyrifera, from reference
collection of Portobello Marine Biological Laboratory
(Dunedin), collected 1.11.1959, from Aquarium Point,
(Portobello, Dunedin, collector unknown. The original
sample consisted of more than one colony, as patches of
female and male gonozooids could be found. Additional
data from label: Hydractinia sp., collected at low tide,
female sporosacs pink, male sporosacs white.

Several living colonies, relatively abundant on Macrocystis
holdfasts and stems of Pyura pachydermata (Ascidiacea) at
low tide around Portobello laboratory, all infertile,
collected May 1994. One clone cultivated for 5 months in
running sea water. Original colony deposited as para type
P-1081.

Description: Stolonal hydroid colonies, with much-
ramified stolons forming a tight meshwork which is
always covered by perisarc. Spines present but rare,
smooth, 0.2-0.3 mm high. Hydranths arise directly
from stolons and have no basal perisarc collar, size of
polyps up to 4 mm when fully grown and expanded,
normally around 2 mm, diameter around 0.3 mm.
Polyps polymorphic with gastrozooids, gonozooids
and tentaculozooids.

Gastrozooids columnar with conical to dome¬
shaped hypostome and one whorl of 12-16 filiform
tentacles of varying length (up to 2 mm). Tentacles
tapering and directed alternately upward, horizontal,
and downward. Hypostome with a conspicuous white
ring of larger microbasic euryteles. Gastrodermal cells
of hydranth body are very large. Colour orange to
pink.

Gonozooids very similar to gastrozooids in size
and shape, only with fewer tentacles (up to 8), also
with broad ring of nematocysts on hypostome. Gono¬
phores are sessile sporosacs arising in the middle
region of the hydranth, 4-6 per hydranth, spherical to
oblong. Females and males in separate colonies.
Females with approximately 14-16 eggs with con¬
spicuous germinal vesicle, spadix present but no radial
or circular canals. Colour, pink. Male gonophores
white, similar to female ones in shape, with spadix,
no radial or circular canals visible, gonads with no

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visible compartmentation. Gonophores bear no
nematocysts.

Few tentaculozooids present, arising directly from
stolons, contracted 1 mm, expanded up to 10 mm,
slightly knobbed at distal end. Nematocysts:

a) microbasic euryteles from hypostome, (11-13) x
(3.5-4.5) Jim, s = 1.

b) microbasic euryteles from tentacles, (9-11.5) x (3.5-
4) Jim.

c) microbasic mastigophores, rare, only frequent in
tentaculozooids, (8-9.5) x (3.5-4.5) Jim, s = 0.8.

d) desmonemes, thread in undischarged capsule with
rope like structure, discharged with 4 to 5 coils,
(5.5-7) x (3-4) Jim.

Additional measurements: stolon diameter 70-80 Jim,
gonophore diameter 0.4-0.5 mm (in preserved
material).

Type Locality: Aquarium Point, near Portobello Marine
Laboratory, Dunedin.

Etymology: The species name otagoensis refers to the

occurrence in the Otago region, South Island.

Remarks: All Stylactaria species differ in minute details
only, but S. otagoensis does not fit any of the other
described species (cf. Namikawa 1991). The most
similar species is 5. conchicola (Yamada, 1947) from
Japan. It is distinct in lacking mastigophores, and in
the substratum, occurring specifically on one gastro¬
pod species only (Namikawa et al 1992a). Other
similar species are S. betkensis Watson, 1978 from
Australia, the widely spread S. inennis (Allman, 1872),
and S. reticulata Hirohito, 1988 from Japan. Stylactaria
betkensis differs from S. otagoensis in its nematocysts
and in having gonophores with radial canals. Sty¬
lactaria inennis also has eumedusoids, and S. reticulata
one egg per gonophore only.

Stolons of S. otagoensis can form a rather tight
mesh work. In fertile colonies, however, the stolons did
not coalesce, and were covered by perisarc, verified
by hand-made thin sections. In culture, no basal-plate
formation (as in Hydractinia) was observed. In fertile

Fig. 30. Stylactaria otagoensis n. sp., from type and living material, a) gastrozooid and tentaculozooid; scale bar 0.5 mm. b)
female gonozooid, same scale as a), c) female gonophore; scale bar 0.1 mm. d) nematocysts in pairs of undischarged and
discharged capsules: microbasic euryteles from hypostome, microbasic euryteles from tentacles, microbasic mastigophores
and desmonemes; scale bar 10 Jim.

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colonies almost all hydranths were gonozooids. This
suggests that there is actually no functional differ¬
entiation into gastro- and gonozooids, but all gastro-
zooids might turn into gonozooids. Reproduction may
be seasonal as no fertile colonies in about six separate
samples could be found in May. Fertile material was
collected in November.

The function of the tentaculozooids has been shown
to be defensive (Namikawa et al. 1992b).

Records from New Zealand: Portobello, Dunedin.

Tiordlandia n.gen.

Stolonal, sessile hydroid colonies arising from ramified
and anastomosing stolons always covered by perisarc.
Polyps dimorphic, with gastrozooids and gonozooids.
Gastrozooids with filiform tentacles scattered in a
band below hypostome. Gonozooids with one whorl
of reduced tentacles and below them gonophores in
several whorls. Gonophores sessile sporosacs. The
hydrorhiza forms erect, branching tubular structures
overarching the gonozooids.

Type Species: Fiordlandia protecta n.sp.

Remarks: The special staghorn-like structures over¬
arching the gonozooids, and the scattered tentacles of
the gastrozooids, make it necessary to create a new
genus. The genus name is derived from the region
where the species was first found: Fiordland, south¬
west South Island. The genus is currently monotypic.

Fiordlandia protecta n.sp. (Figs 31, 32a-f)

Material Examined:

Several well-preserved colonies growing on Symplecto-
scyphus jolmstoni (Hydrozoa, Thecata), collected by M.
Page, 26.11.1993 in Doubtful Sound, 7 m, scuba, preserved
in isopropanol. The Syniplectoscypluis was partially over¬
grown by the bryozoan Fenestrulinn thyreophora and a
foliose red alga; F. protecta was overgrowing both of them;
several colonies present. One part of a male colony was
deposited as holotype H-645, the remainder of the material
was deposited as paratype P-1071 (NIWA).

Description: Epibiontic hydroid colonies growing on
Syrnplectoscyphus jolmstoni, bryozoans, and red algae.
Polyps stolonal, sessile, arising from attached, ramified
stolons. Hydrorhiza covered with perisarc. Polyps
dimorphic with gastropods and gonozooids. Gastro¬
zooids large, up to 3 mm high, club-shaped with
tentacular region swollen, with large dome-shaped
hypostome, 10-20 filiform tentacles of different length

in 3-4 indistinct whorls, or scattered in a band below
hypostome. Tentacles with chordoid gastrodermis.
Gastrozooids more frequently found at periphery of
colony. Gonozooids small, not higher than 1 mm,
mostly below 0.5 mm (preserved material), body
tubular with conical hypostome, with 3-4 tentacles in
one whorl. Tentacles with distal concentration of
nematocysts. Below tentacles 3-10 gonophores in one
to several whorls. Gonophores spherical, up to 250 pm
in diameter, without radial or circular canals, without
tentacles, no or few nematocysts present, with bulbous
spadix and one egg in female, with cylindrical spadix
in male. Male gonad not compartmented. The gono¬
zooids occur in dense aggregations, touching each
other. Between such aggregations of gonozooids,
tubular branching structures like stolons arise and
overarch gonozooids. They have a vertical stem which
then branches horizontally 1-3 times. The horizontal
ends of 2 erect structures can fuse and thus form
bridges. The erect tubes have open distal ends. The
coenosarc reaches to these openings but does not
protrude, and does not contain more nematocysts than
in normal stolons. Nematocysts of both zooids:

a) microbasic euryteles, (8-9.5) x (3.5-4) pm, s = 0.8.

b) desmonemes, discharged with 4-5 coils, (5-
6.4) x (3-4) pm.

Stolon diameter 60-100 pm; protective stolon dia¬
meter: stem 110-160 pm, branches 70-100 pm.
Colours: some orange colour left in the gonophores.

Type Locality: Ranson Head, Crooked Arm, Doubtful
Sound, Fiordland, New Zealand.

Etymology: The species name protecta refers to the
erect branching stolons.

Remarks: The erect, staghorn-like structures (Figs 31,
32c) arising from the hydrorhiza and overarching the
gonozooids are a very special, hitherto unknown
structure. Presumably they are comparable to spines
as found in other Hydractiniidae. These structures are
regarded here as protective stolons. At first, they were
taken as parts of another hydroid, but after dissection
and careful examination of several such elements it
became clear that their coenosarc was continuous
with the remainder of the colony. Their peculiar
branching pattern and their distribution indicated that
they occurred only where the gonozooids grew in
dense aggregations (Fig. 31). They were rare or absent
from the more peripheral parts where more gastro¬
zooids occur. The examined material certainly con¬
tained more than two colonies because female and
male gonozooids were observed in separate patches.
Unfortunately, no other material became available and
therefore it is not known whether such protective

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stolons are always present in mature colonies. The
function of these structures seems quite obvious,
as they can protect the gonophores from mechanical
disturbance and abrasion. For colonies growing in
shallow water on flexible substrata like branched
thecate hydroids this would be advantageous.

The gonophores or even the whole colony may be
intensely coloured when alive as some colour was left
and the fixative (alcohol) was stained dark orange.

Records from New Zealand: Known only from the type
locality.

Fig. 31. Fiordlandia protectan . sp., from preserved material, view of colony; note the characteristic, erect, stolon-like structures;
scale bar 0.5 mm.

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Family RATHKEIDAE Russell, 1953

Colonial hydroids arising from ramified stolons.
Hydranths all alike, very extensible, without a caulus,
with one whorl of filiform tentacles. Gonophores arise
from stolons, or more rarely from base of hydranths.
Gonophores liberated as free medusae. Medusa with
four elongated lips forming oral arms, these either
simple or branched, with terminal knobs, mostly also
with lateral nematocyst knobs. With four or eight
radial canals and a circular canal. Marginal tentacles
in eight groups. Ocelli absent. Gonads encircle manu¬
brium completely (after Bouillon 1985a).

Remarks: Only the genus Rathkea is known from New
Zealand.

Rathkea Brandt, 1838

Polyps with the characteristics of the family. Medusae
with the characteristics of the family but only four
radial canals.

Type Species: Rathkea odopunctata (M. Sars, 1835).

Remarks: The characteristics of all Rathkea species have
been reviewed by O'Sullivan (1984). Only the life cycle
of the type species is known (Rees & Russell 1937).
Two species are known from New Zealand:

R. formosissima : oral arms simple, with one terminal
cluster and several lateral clusters.

R. octopundata : oral arms bifid, each end with a
terminal nematocyst cluster.

Rathkea formosissima (Browne, 1902) (Fig. 33)

Lizzia formosissima Browne, 1902: 278.

Rathkea formosissima : Browne & Kamp 1939: 281, pi. 14, fig.
5, pi. 19, fig. 1; Kramp 1959:104, fig. 76; Kramp 1961: 72;
O r Sullivan 1982: 33, fig. 14, map 13; O'Sullivan 1984: 868;
Barnett 1985: 86, fig. 9B, pi. 3, fig. 3F.

Material Examined:

No material seen.

Fig. 32. Fiordlandia protecta n. sp., from preserved samples, a) gastrozooid, CL; scale bar 0.5 mm. b) male gonozooid with
several gonophores and a concentration of nematocysts in tentacle tips, CL; scale bar 0.5 mm. c) most frequent type of
protective stolon, CL, same scale as in b) d) young male gonophore, CL; scale bar 50 |im, e) female gonophore, same scale as
d). f) nematocysts: microbasic eurytele, same discharged, desmoneme, same discharged; scale bar 10 |im.

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graphs and figure, however, allow a tentative identi¬
fication. The medusae from Leigh were smaller than
1 mm which is about the smallest size documented by
Browne and Kramp (1939). The figures in Barnett
(1985) show medusae with one interradial tentacle
only, which is different form Browne and Kramp
(1939) who observed three tentacles per interradial
bulb. This discrepancy may be explained by the
frequent loss of tentacles during collection, but new
observations should re-investigate this. O'Sullivan
(1984) described Rathkea lizzioides from inland marine
waters of Antarctica. This species is very similar to R.
formosissima and is distinguished from it by characters
normally considered too variable for discrimination
(more oval marginal bulbs, more nematocyst clusters
on oral arms, up to 7 tentacles on the perradial bulbs).
But R. lizzioides belongs to a well-isolated population
and it is at least plausible that it has developed into a
separate species. A re-investigation using allozyme
analysis would be very instructive.

Records from New Zealand: Leigh Marine Reserve
(Barnett 1985).

Fig. 33. Rathkea formosissima , after a photograph of T. Other Records: Stanley Harbour, Falkland Islands.
Barnett; tentacle numbers are uncertain; scale bar 0.2 mm.

Description (after Browne & Kramp 1939): Medusa
with bell-shaped umbrella, slightly higher than wide,
size up to 3 mm, with a dome-shaped apical projection,
lateral walls thin. Manubrium small, quadrangular,
length about one-third of bell cavity, on a broad
peduncle of similar length in older animals, smaller
in younger animals. Mouth with 4 perradial oral arms,
each with 7-11 opposite clusters of nematocysts on its
sides and always with one terminal cluster. Younger
stages with medusae buds on interradial sides of
stomach. Mature animals with gonads on stomach, in
female completely covering stomach, in males divided
by 4 narrow perradial furrows. With 4 narrow radial
canals and 8 tentacular bulbs. Perradial bulbs with 3-
5 tentacles, interradial bulbs with3 tentacles. No ocelli
present.

Polyp stage unknown.

Type Locality: Falkland Islands.

Remarks: Rathkea formosissima was previously known
from its type locality only and nothing is known about
variations in other populations. Barnett (1985)
recorded R. formosissima from Leigh, which is the first
record of this species outside its type locality. She
obtained only five animals in June 1983, none of which
could be re-examined for this study. Barnett's photo-

Rathkea octopimctata (M. Sars, 1835) (Fig. 34a-c)

Cytaeis octopimctata M. Sars, 1835: 28, pi. 6 figs 14a-g.
Rathkea octopimctata : Mayer 1910: 177, pi. 20, fig. 11; Rees &
Russell 1937: 71, figs 7-8; Russell 1953:137, pi. 7, figs 3-
4, text-figs 65A-E, 66, 67A-B; Werner 1958: 138, figs 1-
13; Kramp 1959: 103, fig. 75; Kramp 1961: 72 (cum syn.);
Bouillon & Werner 1965: 137, figs 1-7; Kramp 1968: 30,
fig. 74; Naumov 1969: 212, figs 80-81; Arai &
Brinckmann-Voss 1980: 33, fig. 16; O'Sullivan 1984: 868,
table 2; Werner 1984: fig. 93.

Material Examined:

6 living medusae from surface plankton near Ti Point
(Leigh), collected 7.8.1991, bell ~1 mm high.

10 preserved medusae from Leigh Marine Reserve, collected
by T. Barnett, bell up to 1 mm, some deposited.

Description:

Medusa stage with asexual budding: With spherical
umbrella and a rounded apical projection, bell
diameter around 1 mm. With a gastric peduncle.
Dilated velum spanning half of radius. Manubrium
quadrangular, half as long as bell cavity, mouth margin
with 4 perradial extensions that form oral arms. Bases
of oral arms continued on manubrium for some
distance as perradial ridges. Oral arms distally
branched and both ends with a nematocyst cluster; 1
pair of lateral clusters can be present. Medusae buds
present on stomach. With 4 narrow radial canals and

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b ff\

Fig. 34. Rathkea octopunctata. a) Medusa from Leigh
Marine Reserve; scale bar 0.2 mm. b) nematocysts: desmo-
neme, heteroneme; scale bar 10 mm. c) polyp stage, modified
after Rees and Russell (1937) with permission of Cambridge
University press. Note: the polyp stage is not yet known
from New Zealand; scale bar 0.2 mm.

8 marginal bulbs, 4 perradial, 4 interradial. Each
perradial bulb with 3 tentacles, each interradial bulb
with 2 tentacles. Tentacles thin. No ocelli present.
Nematocysts:

a) heteronemes, (6.5-9) x (2.5-3.5) pm.

b) desmonemes, 4x2 pm.

Mature medusa (after Russell 1953): As above but 3-
4 mm, with up to 5 tentacles per perradial group and

up to 3 per interradial group. Oral arms with 1 or 2
lateral pairs of nematocyst clusters and 1 between the
2 branches. Gonads completely surround the stomach.
Nematocysts:

a) microbasic euryteles, sometimes like mastigo-
phores, in oral arms (10-12) x (3-4) pm.

b) microbasic euryteles, sometimes like mastigo-
phores, in marginal tentacles (6-10.5) x (2-3) pm.

c) desmonemes, in marginal tentacles only, (3-5) x (2-

3) pm.

Polyp stage (after Rees & Russell 1937; Werner 1958):
Hydroid colonies arising from ramified stolons.
Hydranths uniform, up to 0.6 mm high, sessile,
without caulus, base with thin gelatinous perisarc, one
whorl of 4-6 tentacles, hypostome conical. Tentacles
very thin and long. Medusae bud arise from stolons
or rarely from base of hydranth.

Nematocysts (after Bouillon 1985a): microbasic
euryteles, desmonemes.

Type Locality: Norway.

Remarks: The examined Rathkea octopunctata medusae
from New Zealand agree well with other descriptions.
No mature medusae or polyps are so far known from
New Zealand. The polyp has only rarely been reported
from nature, because it is very difficult to find owing
to its small size. Rathkea octopunctata is one of the few
hydromedusae for which a lot of morphological,
physiological, and ecological information is available
(see Kramp 1961; Arai & Brinckmann- Voss 1980, for
references). The life cycle of R. octopunctata is deter¬
mined by seasonal temperature changes. Falling
temperatures in autumn induce the polyps to develop
medusae which then appear in the winter plankton.
The young medusae multiply by budding more
medusae until rising water temperatures in late spring
induce the development of gonads and later sexual
reproduction. The medusae then disappear from the
plankton by late summer (Russell 1953; Werner 1958).
Barnett (1985) observed R. octopunctata near Leigh in
large numbers mostly during winter months. This is
in accordance with observations from the northern
hemisphere.

Records from New Zealand: Leigh Marine Reserve,
Whangateau Harbour (Barnett 1985; this study).

Other Records (medusa): Northwestern Europe; Medi¬
terranean; Black Sea; Barents Sea and White Sea; Ice¬
land; Greenland; Hudson Strait; Newfoundland; New
England; Bermuda; Aleutian Islands; Kamchatka;
northern Japan; British Columbia; Victoria, Australia
(Kramp 1959; Arai & Brinckmann-Voss 1980; O'Sul¬
livan 1984).

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Family CYTAEIDIDAE L. Agassiz, 1862

Hydroids forming nompolymorphic colonies arising
from reticulate stolons covered by perisarc. Hydranths
are without caulus (sessile), with one whorl of filiform
tentacles below conical hypostome. Base of hydranths
often with a perisarc collar. Spines absent. Gonophores
develop from stolons and are liberated as medusa or
remain fixed as sporosacs. Medusa with simple,
circular mouth. With non-branching oral tentacles near
mouth. Gonads develop in interradial position on
manubrium or encircle manubrium. With four, rarely
eight, solid marginal tentacles without lumen. Ocelli
lacking (after Millard 1975; Bouillon 1985a; Calder
1988).

Remarks: Rees (1962) provided a review of this family.
Only Cytaeis is known from New Zealand.

Cytaeis Eschscholtz, 1829

Polyp stage as in family diagnosis. Medusa typical of
Cytaeididae with four marginal tentacles.

Type Species: Cytaeis tetrastyla Eschscholtz, 1829.

Remarks: Only one species is known from the literature
to occur in New Zealand waters.

Cytaeis cf. tetrastyla Eschscholtz, 1829 (Fig. 35a-b)

Cytaeis tetrastyla Eschscholtz, 1829: 104, fig. 2.

Cytaeis tetrastyla: Kramp 1959: 99, fig. 62; Kramp 1961: 63
(cum syn,); Kramp 1968: 26, fig. 64; Brinckmann-Voss
1970: pi. 7, fig. 4; Bouillon 1980.

Material Examined:

No material from the New Zealand region was available; 4
preserved medusae occurred at NZOI Stn A414, 23°16'5,
177°12.5E (south of Fiji Islands), 28.3.1958, surface.

Description: (Material from Stn A414): Medusa bell¬
shaped, widest near base, up to 4 mm high (up to 6
fide Kramp 1961), 3 mm wide. Apical jelly about twice
as thick as lateral walls, without or with very slight
peduncle. Manubrium very voluminous, pear-shaped,
length four-fifths of bell cavity. Manubrium with up
to 20 simple, capitate tentacles scattered in a band near
the mouth, adnate at origin. Stomach base with many
medusae buds. With 4 broad radial canals and thin
circular canal. Epidermis of tentacle bulbs triangular,
flat, attached to exumbrella above tentacles. With 4
tentacles of about bell length when contracted.
Tentacles thick, tapering. Nematocysts:

Fig. 35. Cytaeis cf. tetrastyla from south of Fiji Islands, a)
medusa; scale bar 0.5 mm. b) nematocysts: microbasic
eurytele, same discharged, desmoneme, same discharged,
microbasic mastigophore, same discharged; scale bar 10 pm.

a) microbasic euryteles, (10.5-11.5) x (3.5-4) pm,
s = 0.7.

(b) desmonemes, discharged with 5 coils, (7-10.5) x
(4.5-5.5) pm.

Polyp stage unknown.

Type Locality: First described from the Atlantic Ocean
south of the Equator (Haeckel 1879).

Remarks: There are several Cytaeis species known from
their polyp stage only and many of them seem to pro¬
duce similar medusae. It may therefore be that the

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current concept of Cytaeis tetrastyla includes more than
one species (cf. Rees 1962; Pages et al 1992). Therefore
C. tetrastyla was here identified only tentatively.
Kramp (1965) recorded C. tetrastyla near the Kermadec
Islands. Probably it does not occur near the main
islands of New Zealand, being a warm-water species
distributed between 40°N and 30°S (Kramp 1965).

Records from New Zealand: Kermadec Trench (Kramp
1965).

Distribution: Tropical to subtropical seas (Kramp 1965;
van der Spoel & Bleeker 1988).

Family PANDEIDAE Haeckel, 1879

Hydroid stage normally stolonal, not branching, with
spindle-shaped hydranths bearing one whorl of
filiform tentacles (exceptionally two whorls, scattered
tentacles or no tentacles). Perisarc developed to a
variable degree, occasionally missing completely, but
often also enveloping hydranth as a filmy pseudo¬
hydrotheca. Reproduction is by free medusae (except
in some genera of questionable affinity). Medusae with
or without an apical projection. Manubrium large,
mounted or not on a peduncle; mouth with either four
simple, or crenulated, or complexly folded lips. With
four (rarely eight as in Octotiara) radial canals, often
broadened to a band; exceptionally centripetal canals
can be present. Mesenteries are frequent. Gonads
either with a smooth surface or complexly folded; they
differentiate on the manubrium in adradial or inter-
radial positions; in a few species they extend onto the
radial canals. Tentacles hollow, originating from
conical tentacle bulbs, tapering, often laterally com¬
pressed. The tentacles never have terminal nemato-
cyst clusters (capitation). There may be rudimentary
tentacles, cirri-like tentacles, or marginal protuber¬
ances. Ocelli may be present or absent (after Bouillon
1985a).

Remarks: The above definition is rather vague and com¬
plex. It is probable that this group is not monophyletic
and future revisions will become necessary (see also
under families Clavidae and Bougainvilliidae). A
synapomorphy for this family may be the presence
of only two opposite tentacles in newly released
medusae. However, at present only a fraction of the
life cycles are known and it is premature to promote
this character as a synapomorphy. Characteristics of
genera known from New Zealand:

Amphinema : adult medusae with two tentacles, polyps
bend head when stressed.

Annatiara : medusa with nematocyst tracks on exum¬

brella, without apical process, with broad, cruci¬
form stomach.

Halitliolus : medusae with four to eight tentacles that
are not laterally compressed, without mesenteries.
Leuckartiara : medusae with laterally compressed
tentacles, with mesenteries between radial canals
and manubrium, gonad folds directed towards
perradial.

Neoturris : medusa with gonad folds directed towards
interradial, with mesenteries, small apical process,
tentacles compressed laterally.

Pandea : medusae with reticulate gonads, with exum-
brellar nematocyst tracks, tentacles compressed
laterally, polyps on pteropod snails.

Pandeopsis : medusa with broad, quadrangular
stomach, smooth gonads, with mesenteries, with¬
out cirri.

Merga: medusa with cruciform base of stomach,
smooth gonads, more than four fully developed
tentacles, without cirri-like tentacles, with
mesenteries.

Barnettia ; medusa with smooth gonads, eight fully
developed tentacles, cirri-like small tentacles.

Amphinema Haeckel, 1879

Stolonal hydroid colonies, hydranths with a well-
developed caulus, caulus longer than hydranth,
covered by perisarc. Hydranths without a pseudo¬
hydrotheca, spindle-shaped, with one whorl of filiform
tentacles, with conical hypostome. Gonophores arise
either from cauli, stolons or both. Gonophores are
released as free medusae. Medusae with two tentacles
in opposite position. Gastric peduncule absent.
Generally with a considerable apical projection. There
may be marginal cirri or protuberances. With or
without mesenteries. With or without ocelli. Gonads
on manubrium in adradial or interradial position,
occasionally extending to the radial canals.

Type Species: Amphinema dinema (Peron & Lesueur,
1809).

Remarks: The above diagnosis was slightly modified
after Bouillon (1985a). Amphinema polyps are rather
inconspicuous and very similar to other polyps in the
Pandeidae and Bougainvilliidae. But many Amphinema
polyps, if not most, show a characteristic reaction to
stimuli: they bend back their hydranths so that the
mouths face towards the substratum (cf. also Boero &
Bouillon 1989). This reaction was seen in both species
occurring in New Zealand (Figs 36a, 37a) and differs
from other similar polyps that retract or contract upon
disturbance. The gastrodermis of the hydranth body

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is composed of large, vacuolated cells. Although not

diagnostic, it can help to identify Amphinema polyps.

Characteristics of species known from New Zealand:

Amphinema dinema : Medusae buds on short stalks on
stolons only. Medusae without folds in gonads,
green manubrium, no short tentacles between long
ones.

Amphinema rugosum: Medusae buds on short stalks on
stolons and cauli. Medusae with gonads in three
folds, yellow manubrium, short tentacles between
long pair.

Amphinema dinema (Peron & Lesueur, 1809)

(Fig. 36a-d)

Oceania dinema Peron & Lesueur, 1809: 346.

Perigonimits serpens Allman, 1863: 10.

Stomotoca dinema: Mayer, 1910: 109, pi. 9, figs 8-10, pi. 10,
figs 1-4.

Amphinema dinema : Rees & Russell 1937: 62, figs 1-4; Russell
1953: 180, pi. 10, figs 1, 2, 4, pi. 11, figs 1, 3, text-fig. 89
(cum syn.); Kramp 1959: 117, fig. 109; Kramp 1961: 93;
Kramp 1968: 42, fig. 108; Goy et al 1991: 109, fig.24.

Material Examined:

Fertile colony collected beneath Queens Wharf, Wellington
Harbour, 25.1.1994, subtidal, growing mainly on Water-
sipora subtorquata (Bryozoa) and sponges; more than 100
hydranths with at least 30 medusae buds; part of colony
deposited.

Medusae released from above colony grown to maturity,
material deposited.

Several mature medusae collected from plankton at Sea-
toun, Wellington Harbour, some deposited.

Description:

Polyp stage: Stolonal hydroid colonies arising from
attached, ramified stolons. Cauli not branched. Polyps
up to 1.5 mm high, with cauli covered by perisarc.
Perisarc at proximal origin only occasionally annu-
lated, becoming very thin distally and the distal
perisarc margin on hydranth body difficult to observe.
Perisarc can be infested with detritus. Hydranth
spindle-shaped with a dome-shaped hypostome.
Below hypostome one whorl of 8-11 filiform tentacles
alternately pointing up- and downwards. Medusae
buds arise from stolons only, with stems shorter than

Fig. 36. Amphinema dinema all from life, a) hydroid colony with medusae buds; scale bar 0.5 mm. b) newly hatched medusa;
scale bar 0.2 mm. c) nematocysts of medusa: microbasic euryteles; scale bar 10 pm. d) mature medusa grown from polyp,
identical animals were also found in the plankton, gonads shaded; scale bar 0.5 mm.

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bud height, Nematocysts and additional observations:

a) microbasic euryteles, (7.5-8) x (2-2.5) mm, s = 1.1.

b) desmonemes, discharged with 3 coils, (4-4.5) x (2-
2.5) mm.

Colour of hydranths and medusae buds intense red-
orange; medusa bud height 280-340 mm, stems of
medusae buds 55-60 mm, caulus diameter proximally
50-75 mm, distally 120 mm.

Newly hatched medusa and juvenile stages: Newly
released medusa 0.6 mm high and 0.5 mm in diameter,
jelly thin, without apical process, exumbrella with
scattered nematocysts. Velum spanning three-quarters
of radius. Manubrium length half of bell height,
conical, red in colour. With 4 radial canals and circular
canal One pair of opposite tentacles, with broad base,
then rapidly tapering. The other 2 radial canals end in
a very reduced tentacle bulb.

Interradial rudimentary bulbs may grow after a few
days; an apical projection starts to grow as well, the
manubrium develops an intensive green colour,
becoming cruciform, tentacles turn red or purple.

Mature medusa (animals from plankton and culture):
Up to 2.5 mm inclusive of large apical projection, bell
slightly wider than high, jelly of uniform thickness
besides top. With slight perradial furrows in top of
umbrella. Dilated velum spanning half of radius.
Manubrium with intensive green colour, manubrium
length three-quarters of subumbrella, cruciform in
section, mouth with 4 prominent, slightly recurved
lips. Eight gonads in adradial pairs, with smooth
surface. With 4 broad radial canals. Two diametrically
opposed tentacles with thick base, tapering and very
long (10 times bell size), deep purple to violet colour;
additionally ca. 12 marginal warts, but no short ten¬
tacles. Ocelli not present. Nematocysts and additional
observations:

a) only microbasic euryteles, (6.5-7) x (2-2.5) pm.
Size of spawned eggs 173 pm (n = 5, s.d. = 12.5 pm).

Remarks on Variation: According to Russell (1953) the
medusa can attain 6 mm in height. Shape and size of
the apical projection of the medusa vary considerably.

General Remarks: It was possible to grow medusae
liberated from polyp colonies to full maturity (spawn¬
ing) (Fig. 36d). The cultivated medusae corresponded
exactly to those found in the plankton and also to
descriptions in the literature. The presence of A. dinemn
has not been noted in New Zealand before. It is
possible that it is a recently introduced species.

Records from New Zealand: Wellington Harbour (new
record of polyp and medusa).

Other Records: NE Australia; Vietnam; India; Medi¬
terranean; Gulf of Guinea; NW Europe; North America
from Florida to Cape Cod (Kramp 1968).

Amphinema rugosum (Mayer, 1900a) (Fig. 37a-d)

Stomotocn rugosn Mayer, 1900a: 4, pi. 2, fig. 5.

Stomntocn rugosn : Uchida 1927: 202, fig. 32; Ralph 1953: 74,
fig. 17.

Amphinema rugosum : Rees & Russell 1937:67, figs 5-6; Russell
1953: 183, pi. 10, fig. 3, pi. 11, figs 2 & 4, text-figs 90A-B;
Kramp 1959: 117, fig. 110; Kramp 1961: 94 (cum syn.);
Kramp 1968: 43, fig. 110; Roper et nl 1983: table 2;
Bouillon 1995: 224.

Material Examined:

Polyp colony on tube of serpulid polychaete, collected
28.2.1994 near Greta Point, subtidal, with many medusae
buds, medusae released and cultivated for 9 days, lost by
accident before reaching maturity, some polyp material
deposited.

Living medusae from plankton near Seatoun, Wellington,
15.4.1994, several juvenile and one mature female.

Ca. 5 medusae collected by T. Barnett, Leigh Marine Reserve,
Leigh, 1983-1984.

Description:

Polyp stage: Stolonal hydroid colonies arising from
attached, ramified stolons. Polyps up to 3.5 mm high,
on cauli up to 2.5 mm long covered by perisarc. Peri-
sarc at proximal origin annulated in all polyps,
expanding at distal end. Perisarc ends in the middle
of the hydranth body, where it adheres to polyp with
a well-marked end. The perisarc can be infested with
detritus. Hydranth around 1 mm high, spindle- shaped
with a dome-shaped hypostome. Below hypostome
one whorl of 10-12 filiform tentacles reaching a length
of 1 mm, alternately pointing up- and downwards.
Medusae buds arise from stolons and from cauli, with
stems shorter than bud height, up to 3 buds per caulus,
Nematocysts and additional observations:

Ocelli not present. Nematocysts:

a) microbasic euryteles, (9-10) x (3) pm, s = 1.3.

b) desmonemes, (5) x (3) pm.

Colour of hydranth white to slightly orange, stems of
medusae buds 100 pm, caulus diameter proximally
80-100 pm, distally 200 pm.

Newly hatched medusa and juvenile stages: Newly
released medusa 0.6 mm high and 0.5 mm in diameter,
jelly thin, a little apical process is mostly present,
exumbrella with scattered nematocysts. Manubrium
length half of bell height, conical, yellow colour. With
4 radial canals and circular canal. One pair of opposite
tentacles with broad base then rapidly tapering, yellow
colour; the other 2 radial canals end in small tentacle
bulbs.

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After a few days short tentacles (50 pm) grow from
the rudimentary bulbs and in the 4 interradial
positions; apical projection grows as well, manubrium
remains yellow-brown.

Mature medusa (from plankton: Up to 4 mm inclusive
of large apical projection, bell slightly higher than
wide, jelly uniformly thin besides top. With slight
perradial furrows in top umbrella. Manubrium yellow-
coloured, reaching almost to velum, cruciform in
section; mouth with 4 prominent, slightly recurved
lips. Eight gonads in adradial pairs, with 3 character¬
istic folds directed interradially. Four broad radial
canals with jagged and smooth margins. Two dia¬
metrically opposed tentacles with thick base, taper¬
ing and very long (ten times bell size), yellow colour.

Additionally ca. 14 shorter (one-tenth bell length)
reduced tentacles. Ocelli not present. Nematocysts:

a) microbasic euryteles, (9-10) x (3) pm, s = 1.3.

b) desmonemes, (5) x (30 pm

Type Locality: First described from Atlantic coast of
the USA (Mayer 1900a).

Remarks: According to Russell (1953) the medusa can
reach 5 to 6 mm in height, and there may be up to 24
small tentacles. Although the New Zealand medusae
released from the polyps could not be grown to full
maturity, the polyp can reliably be referred to A.
rugosinn as it matched very well the description given
by Rees and Russell (1937). The apical process of the
newly-hatched medusa, the yellow colour, and the

Fig. 37. Amphinema rugosinn, all from life, a) hydroid colony with medusae buds; hydranth at right shows the typical bend-
back reaction; scale bar 0.5 mm. b) newly liberated medusa; scale bar 0.2 mm. c) nematocysts of polyp and medusa, micro-
basic euryteles and desmonemes; scale bar 10 pm. d) mature medusa from plankton, gonads shaded; scale bar 0.5 mm.

6 o

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short tentacles of the juveniles are also in accordance
with the descriptions of Rees and Russell (1937). The
adult medusae from the plankton near the same
locality also agreed well with other descriptions.

Records from New Zealand: Polyp: Wellington Har¬
bour (first record by this study). Medusa: Cook Strait
(Kaberry 1937; Ralph 1953, as Stonwtocci rugosa); off
Chatham Islands (Kramp 1965); Avon-Heathcote
estuary in Christchurch (Roper et al 1983); Goat
Island, Leigh (Barnett 1985); Coromandel, Cook Strait
(Bouillon 1995); Wellington Harbour (this study).

Other Records: New Caledonia; Sumatra; China Sea;
Madagascar; Caribbean; Florida to Cape Cod; Adriatic
Sea; north western Europe (Kramp 1968).

Annatiara Russell, 1940

Pandeid medusae with umbrella lacking an apical
process, with exumbrellar nematocyst tracks. Broad
manubrium, not extending beyond umbrella margin,
mouth with much crenulated lips. With four radial
canals. Gonads as folds on interradial sides of stomach.
Many marginal tentacles with elongated, laterally
compressed conical bases, without well developed
abaxial spurs. Rudimentary tentacles present, without
marginal warts or marginal tentaculae. Each tentacle
with an abaxial ocellus.

Type Species: Annatiara affinis (Hartlaub, 1914).

Remarks: No polyp stages are known in this genus.
Contrary to the descriptions of other authors (e.g.,
Kramp 1959, 1961, 1968; Bouillon 1985a) the medusa
of Annatiara affinis has ocelli (Russell 1953; Schuchert
pers. obs.). Only one species is known from New
Zealand.

Annatiara affinis (Hartlaub, 1914) (Fig. 38)

Tinrannn affinis Hartlaub, 1914: 269, figs 221-222.

Tiaranna affinis: Kramp 1926: 68, pi. 1, figs 156-17, map 11.
Annatiara affinis: Russell 1953: 200, figs 101-103; Kramp 1959:
122, fig. 123; Kramp 1961: 96; Kramp 1965: 34; Kramp
1968: 48, fig. 125; O'Sullivan 1982: 41, fig. 18, map 17;
Bleeker & van der Spoel 1988:231, figs 10-12, map; Bouil¬
lon 1995: 224.

Material Examined:

1 medusa from Dana Stn3631II, 18.12.1928, 35°40S, 176°40'E,
1000 m wire, very damaged.

1 medusa from Dana Stn 3630V, 17.12.1928, 34°24'S,
178°42.5'E, 600 m wire, very damaged.

1 medusa from Dana Stn 3626VIII, 13.12.1928, 27°00'S,

177°4TW, 1500 m wire, very damaged.

1 medusa from NZOI Stn N371,10.12.1974,34°23'S, 171°55'E,
0-200 m, damaged, 12 mm, yellow gonads as folds and
pits present, margin very damaged.

1 medusa from NZOI Stn U799, 7.8.1990,42°33.8’S, 170 o 34'E,
depth of haul not known, 10 mm high, some tentacles well
preserved, with abaxial ocelli.

Description: Medusa with bell-shaped umbrella, up
to 12 mm high and 15 mm wide, without apical pro¬
jection, jelly uniformly thick, thickness moderate. Ex¬
umbrella with meridional tracks of nematocysts that
originate from each marginal bulb. Manubrium short
but very broad, cross-shaped, attached to subumbrella
along the 4 perradial arms of the cross. Walls of manu¬
brium rather thin. Mouth with crenulated margins.
Gonads in irregular fold and pits along perradial side
of stomach. Four broad radial canals and circular
canal; 16 full tentacles and 16 short rudimentary ten¬
tacles; all tentacles have a laterally compressed base
that clasps the umbrella margin, all with an abaxial
brown ocellus. Large tentacles rather short, with broad
elongated base, then tapering.

Type Locality: 48°29 r N, 13°55'W (southwest of Ire¬
land).

Remarks: The diameter of the bell may reach 23 mm
with up to 44 large tentacles (Russell 1953).

Fig. 38. Annatiara affinis, after preserved sample from Stn
U799, posterior tentacles not shown, some tentacles recon¬
structed; scale bar 1 mm.

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Records from New Zealand: East of North Island
(Kramp 1965), SW of Three King Islands (Bouillon
1995), off Greymouth (Westland) (this study).

Other Records: Atlantic Ocean between 60°N and the
equator; Southwest Africa; central Indian Ocean; Sri
Lanka; East of Australia; Indonesia; Papua New
Guinea; generally in deeper waters (Kramp 1965;
Bouillon 1980; Bleeker & van der Spoel 1988).

Halitholus Hartlaub, 1913

Pandeid medusae bearing a large, dome-shaped apical
projection, with quadrangular manubrium, with
gonad folds directed outwardly. Mouth margin crenu-
lated. Mesenteries absent. With four or more tentacles.

Type Species: Halitholus pauper Hartlaub, 1913.

Remarks: Probably only the life cycle of Halitholus
cirratus Hartlaub, 1913 is known (Naumov 1969, as
Perigonimus yoldia-arcticae), although it is not clear
whether this life cycle was really revealed through
rearing experiments. Rees and Thursfield (1965)
attributed a "Perigonimus 1 ' type of hydroid to the
medusa Halitholus intermedins. This was solely based
on the occurrence at the same locality (Stanley Har¬
bour, Falkland Islands). Although the hydroid might
well have been H. intermedius, the uncertainty is too
great and the life cycle of H. intermedius is better
regarded as still unknown.

As noted by Arai and Brinckmann-Voss (1980),
Halitholus species are often difficult to distinguish from
Leuckartiara species, especially when immature.

Only one Halitholus species is known from New
Zealand.

Halitholus pauper Hartlaub, 1913 (Fig. 39)

Halitholus pauper: Hartlaub 1913: 272, figs 223-224; Kramp
1926: 71, pi. 2, figs 1-3; Kramp 1959: 119, fig. 116; Kramp
1961: 111; Arai & Brinckmann-Voss 1980: 46, fig. 22.

Material Examined:

3 medusae, size approx. 5 mm, but shrunken, from MoNZ,
in two tubes, apparently females as eggs visible. Label tube
1: "Zool. Dep. Victoria University Wellington, Leuckartiara
octona, WHP." Label tube 2: "N.Z. Oceanographic Institute
Wellington, Leuckartiara octona, Wgtn Hbr, Dec 1962."

Description:

Medusa ca. 5 mm high, bell-shaped, with low, rounded
apical process, lateral walls thin. Manubrium massive,
reaching two-thirds of subumbrellar height, cross¬
shaped in section. Mouth cruciform, margin moder¬

ately crenulated. Four gonads on interradial sides of
stomach in the form of a horseshoe, opening directed
downwards. Gonads with folds, the openings directed
perradially and upwards. Four radial canals, rather
broad, margins partly smooth, partly jagged. No
mesenteries. Ring canal smooth. Four perradial ten¬
tacles, with thick base, then tapering, not laterally
compressed, clasping bell margin. Four larger inter
radial bulbs and 8 smaller adradial bulbs, all without
tentacles. All bulbs with an small, abaxial, red ocellus.

Type Locality: Western Greenland.

Remarks: The present samples closely resemble
Hartlaub's (1913) figures of Halitholus pauper with 4
tentacles (although the tentacle bases in Hartlaub's
figure are drawn incorrectly as noted by Kramp 1926).
In the formal diagnosis given by Kramp (1959, 1961,

Fig. 39. Halitholus pauper, from Wellington Harbour; scale
bar 1 mm.

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1968), there should be 4 interradial tentacles of smaller
size. According to Kramp (1926) these do not develop
until the medusa becomes 5 mm high. At this size the
gonads are already fully developed (Hartlaub 1913).
The description of the gonad as horse-shoe shaped was
taken from Hartlaub (1913) and has also been used by
later authors. The shape of the gonads of the present
material corresponds with Hartlaub's figures, but they
are better described as pairs of adradial gonads con¬
nected by a transverse interradial bridge. This inter¬
radial connection was found in the majority of the
gonads observed in the present study. With hesitation,
the medusae from Wellington were therefore assigned
to H. pauper. Some caution must be stated because Arai
and Brickmann-Voss (1980) noted that there may be
more unknown Halitholus species that are difficult to
separate.

It is rather surprising to find H. pauper in the south¬
ern hemisphere, where one would rather expect to find
H. intermedius (Browne, 1902), known from the
Falkland Islands and South Africa (Kramp 1961). This
species is not easily separated from H. pauper . The
main distinctions are: pairs of adradial gonads or a
horse-shoe shape that is merely indicated, a large,
pointed apical projection, and larger ocelli (Browne &
Kramp 1939).

Records from New Zealand: Wellington Harbour (new
record).

Other Records: Arctic parts of Atlantic; Iceland, Puget
Sound (Canada); Japan.

Leuckartiara Hartlaub, 1913

Hydroids colonial, arising from stolons. Hydrocauli
not or sparingly branched, covered by perisarc and
not fascicled. Perisarc extends onto hydranth body as
a more or less gelatinous pseudohydrotheca which
does not envelop the tentacles. Hydranths with a
conical hypostome and one whorl of filiform tentacles.
Gonophores develop on cauli or hydrorhiza and are
covered by thin perisarc, liberated as free medusae.
Medusa typically pandeid, mostly with an apical
projection of variable size. Manubrium voluminous,
connected to radial canals by mesenteries. Mouth with
extensively folded and crenulated margin. Gonads on
interradial walls of manubrium in the shape of a horse¬
shoe, with folds directed towards outside. Radial canal
very broad, often jagged. With many tentacles arising
from elongated bulbs, laterally compressed, rudi¬
mentary tentacles often present. With or without ocelli
(Bouillon 1985a).

Type Species: Leuckartiara octona (Fleming, 1823).

Remarks: Younger specimens of Leuckartiara may lack
mesenteries and are often difficult to distinguish from
Halitholus species. Only one species of Leuckartiara is
known from New Zealand with certainty, more may
occur (see below).

Leuckartiara octona (Fleming, 1823) (Fig. 40a-b)
Geryonia octona Fleming, 1823: 299.

Leuckartiara octona: Hartlaub 1914:285, figs 238-214, figs 244-
253; Uchida 1927: 211, fig. 37; Rees 1938: 11, figs 3-5;
Russell 1953: 188, pi. 11, figs 5-6, pi, 12, fig. 3, pl2 30-31,
text-figs 91-96; Kramp 1959: 121, fig. 119; Kramp 1961:
105 (cum syn); Kramp 1968: 47, fig. 121; Millard 1975:
123, figs 4la-d; Arai & Brinckmann-Voss 1980: fig. 29;
Pages et al. 1992: 10, fig. 11; Bouillon 1995: 224.

Material Examined:

1 medusa from NZOI Stn N356, 36°31.3 r S, 175°17.6 r E
(Hauraki Gulf), depth 0-50 m, with 14 tentacles, 6 mm
bell and 2 mm process.

1 medusa collected by T. Barnett, 1983, Leigh Marine
Reserve, 6 mm high, 8 tentacles, damaged, mesenteries
not seen, identification uncertain.

Description:

Medusa stage: Adult up to 8 mm high (up to 20 mm
from literature), higher than wide, with conical or
spherical apical projection. Lateral walls thin. Manu¬
brium large, length up to two-thirds of bell cavity,
connected to radial canals by mesenteries for half of
its length. Mouth large, with much folded margin. Four
gonads covering whole interradial area, bipartite but
connected interradially, with many folds directed
towards perradial. Four very broad radial canals and
a much thinner circular canal; radial canals often with
jagged margins. Up to 14 (up to 24 from literature)
long tentacles. Tentacles with a long conical base,
laterally compressed, clasping umbrella margin and
forming an abaxial spur which bears an ocellus. In
between each pair of long tentacles a rudimentary
tentacle bulb.

Nematocyst (after Russell 1938):

a) microbasic euryteles (8-10.5) x (3-3.5) pm.

b) mastigophore (7) x (3.5-4) pm.

Polyp stage (after Millard 1975, not known from NZ):
Hydroid colonies reaching a maximum height of
5 mm. Stems increasing in diameter from base dis-
tally, bearing a terminal hydranth and occasionally 1-
3 laterally. Perisarc firm, often annulated or wrinkled,
especially at base, expanding to form a gelatinous
pseudohydrotheca over base of hydranth, usually
covered with adherent silt. Hydranth with 6-12 ten-

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Fig. 40. Leuckartiara octona. a) preserved from Stn N356;
scale bar 1 mm. b) polyp colony with medusae buds,
redrawn from Rees (1938) with permission of Cambridge
University Press. Maximal height 2.5 mm. Note: the polyp
stage of L octonn has not yet been found in New Zealand.

tacles. Medusa buds with short stems on cauli of
hydranths, rarely on hydrorhiza, reaching a diameter
of 0.45 mm. Newly released medusa with 2 opposite
marginal tentacles. Nematocysts (after Bouillon 1985a):
desmonemes and microbasic euryteles.

Type Locality: Bell Rock, Scotland.

Remarks: The life cycle of Leuckartiara octona was
revealed by Rees (1938), who also gave additional
numeric data. Although not fully grown, the medusa
from Stn. N356 agreed rather well with the existing
description and figures. The first record of this species
for New Zealand was by Kaberry (1937, as Tunis
vesicaria ). He found his animals in the plankton near
Wellington during late autumn. His detailed
description and figure leave no doubt that he actually
had L. octona . Kaberry's medusae reached the size of
25 mm and had up to 24 long tentacles and as many
rudimentary ones, therefore appearing much more like
the forms known from Europe. Ralph (1953, fig. 19)
copied Kaberry's figure and incorrectly named it
Neoturris vesicaria .

The medusae found by Barnett (1985) were very
small and did not have more than 8 tentacles. This
identification is therefore somewhat uncertain as her
medusae may belong to a different species.

The polyp of L. octona is not known from New
Zealand, but should occur here. It often occurs
epizoically on gastropod shells (Rees 1938a; Millard
1975) or on parapodia of the polychaete Aphrodite
(Latham 1963; Schuchert pers. obs.)

Records from New Zealand: Cook Strait (Kaberry
1937), east of Kaikoura (Kramp 1965), ? Leigh Marine
Reserve (Barnett 1985), Hauraki Gulf (Bouillon 1995).

Other Records: Polyp: England; South Africa (Millard
1975). Medusa: European coasts from Portugal to
Lofoten; Iceland; North America from Labrador to
Cape Cod; Mediterranean; west Coast of Africa; Tris¬
tan du Cunha; India; Malayan Archipelago; north¬
eastern Australia; Low Archipelago; China; Japan;
Vancouver; Chile (Kramp 1959).

Neoturris Hartlaub, 1913

Colonial hydroids arising from stolons, stems not
branching. Hydranths on simple cauli, these covered
by perisarc that continues up to the hydranth body as
a pseudohydrotheca but does not envelop the ten¬
tacles. One whorl of filiform tentacles. Gonophores
developing from hydrocauli, sometimes from stolons.
Gonophores completely covered by thin perisarc and

@(D®(|

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liberated as free medusae. Pandeid medusae with an
apical process of variable size, often reduced. With a
very voluminous manubrium connected to radial
canals by mesenteries. Eight gonads in adradial
position on stomach, with folds directed towards
interradial. Interradial part of stomach may have
additional isolated pits of gonads. Eight or more
hollow marginal tentacles with a laterally compressed
base. No marginal warts . Mostly without ocelli (after
Bouillon 1985a).

Type Species: hJeoturris pileata (Forsskal, 1775).

Remarks: Only the life cycle of the type species is known
(Edwards 1965). Neoturris may appear similar to
Leuckartiara, but both genera are readily distinguished
by the direction of the gonad folds. Only N. papua is
known from New Zealand.

NeoUirris papua (Lesson, 1843) (Fig. 41)

Turns pnpun Lesson, 1843: 283.

Neoturris pnpun : Uchida 1927: 210, fig. 36; Ranson 1929, 209,
figure; Kramp 1961: 108; Kramp 1968: 49, fig. 126; van
der Spoel & Bleeker 1988:167, figs 11-12; Bouillon 1995:
229, fig. 4.

Material Examined:

1 medusa from NZOI Stn N400, 40°26.8'S, 175°9.2'E (Cook
Strait, near Levin), 16.12.1974, 0-53 m, well-preserved
female, 11 mm high, 12 tentacles, no ocelli visible.

Description: Large medusa, 11 mm high (may be up
to 18 mm high from literature), much higher than wide,
conical bell, with small apical projection containing
an apical canal. Bell wall uniformly thin. Exumbrella
with longitudinal ridges containing nematocysts. As
many ridges as normal tentacles, ridges are a con¬
tinuation of the abaxial spurs of the tentacle bulbs.
Manubrium very voluminous, two-thirds as long as
bell cavity, connected to radial canals by long mesen¬
teries. Mesenteries as long as half bell height. Lips
extraordinarily complexly folded. Eight gonads in
adradial position on manubrium, covering three-
quarters of its length. Interradial position of manu¬
brium free of gonads. Gonads with a series of hori¬
zontal folds which are directed towards interradial.
Four very broad radial canals and a much narrower
circular canal. Radial canals with smooth margins. Up
to 12 large marginal tentacles, the perradial ones
longer than the others. Tentacle bulbs long, conical,
laterally compressed and clasping umbrella margin.
Between each pair of these marginal tentacles 3 thin
and short tentaculae with reduced bulbs.

Polyp stage unknown.

Fig. 41. Neoturris papua from Stn N400, rear tentacles not
shown; scale bar 1 mm.

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Remarks: All normal tentacles and also the tentaculae
have an abaxial ocellus on their bulbs (Bouillon 1995).

Type Locality: Originally described as Tunis papua
from Waigiou Island, Indonesia.

Records from New Zealand: Greater Cook Strait, off
Horowhenua coast (Bouillon 1995).

Distribution: Widely distributed in the coastal waters
of the tropical parts of the Indo-West-Pacific region
(Kramp 1968).

Pandea Lesson, 1843

Hydroids, where known, forming stolonial colonies
with naked hydranths on a very short caulus.
Hydranth with filiform tentacles. Medusae buds arise
from hydrorhiza. Medusae with the characteristics of
Pandeidae, with or without apical projection, with or
without longitudinal ridges on exumbrella. Lips large
and folded. Gonads at first in the adradii and event¬
ually covering the manubrium, forming a complex
reticulated network. Radial canals band-like. More
than eight tentacles.

Type Species: Pandea conica (Quoy & Gaimard, 1827).

Remarks: So far only the life cycle of P. conica is known
(Picard 1956). Only this species is known from New
Zealand.

Pandea conica (Quoy & Gaimard, 1827)

(Fig. 42a-c)

Dimmed conica Quoy & Gaimard, 1827:182, pi. 6A, figs 3-4.

Pandea conica : Mayer 1910: 118, fig. 63; Uchida 1927: 214,
fig. 38; Russell 1953: 207, figs 107-110; Picard 1956: 1,
figs 1-3; Kramp 1959:123, fig. 127; Kramp 1961: 111 (cum
si/;?.); Kramp 1968: 51, fig. 134; Brinckmann-Voss 1970:
pi. 11, fig. 2; Lalli & Gilmer 1989: 127, fig. 47b; Pages et
nl 1992: 14, fig. 13; Bouillon 1995: 224.

Material Examined:

1 medusa from NZOI Stn N449, 44°28.9'S, 167 # 38.6'E,
1.2.1975, 0-200 m.

3 medusae from NZOI Stn X480F, 41*20.45, 179°05.8“E,
17.10.1993, 25-50 m.

1 medusa from NZOI Stn X480F, 200-400 m.

Description:

Medusa stage: Adult up to 21 mm high, higher than

wide, lateral walls oblique, rather straight, with a small

conical process at apex the tip of which is covered by

opaque epidermis. Exumbrella with many longi¬
tudinal nematocyst tracks that originate from each
tentacle. Jelly moderately thick, thicker at top. Manu¬
brium very large, up to two-thirds of height of bell
cavity. Mouth with four perradial lips with folded or
crenulated margin. Manubrium connected to radial
canals by long mesenteries. Gonads large, filling entire
interradial walls of stomach, forming a coarse-meshed
network of ridges with pits between. Four broad radial
canals with jagged margins, circular canal narrower
and smooth. Up to 24 smooth, hollow, marginal ten
tacles, each with conical, laterally compressed bulb
clasping the umbrella margin. Each tentacle with an
abaxial ocellus. Lacking rudimentary tentacles.
Nematocysts:

a) basitrichous isorhizas (?), only few present, (20-
21) x (19-21) pm.

b) microbasic euryteles, (9.5-11) x (4.5-5) pm, s ~ 0.9.

c) atrichous isorhiza, diameter 5 pm.

Polyp and young medusa (after Picard 1956): Colonial
hydroids living on the planktonic gastropod Clio
cuspidata, arising from attached, ramified stolons.
Stolons fine, covered by thin perisarc. Polyps with a
very short caulus, almost sessile. Hypostome conical.
Eight filiform tentacles of variable length in 2 closely-
set whorls. Medusae buds arise directly from stolons
and have a short caulus, buds covered with thin
perisarc. Newly released medusa as high as wide, bell¬
shaped with moderately thick jelly. Jelly at apex
interrupted by remains of the connection with the
caulus. Manubrium length half of bell cavity, conical
with4 very inconspicuous lips, no gonads visible. Four
broad, smooth radial canals and circular canal. Two
opposite larger and two smaller tentacle bulbs present-
only larger bulbs bear a tentacle. Ocelli not present,
but a dispersed red pigment present in bulbs. Above
bulbs on exumbrella patches of tissue with
nematocysts that later grow towards top. Nematocysts
of polyp:

a) microbasic euryteles, (8-10) x (3.5-5) pm.

b) desmonemes, 5 x 3.5 pm.

Type Locality: Gibraltar.

Remarks: According to Russell (1953) the manubrium
may be reddish, brownish, or yellowish and the size
of the bell can attain 30 mm. The polyps of this species
occur on the shell of the euthecosome pteropod Clio
cuspidata (see Picard 1956, Lalli & Gilmer 1989). New
Zealand lies within the high abundance area of C.
cuspidata (van der Spoel & ETeyman 1983, fig. 48) and
therefore the polyps of P. conica might be expected
here.

The nematocysts found in the medusae from New

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Zealand were more varied than reported by Picard
(1956). He did not mention the large capsule, probably
an isorhiza, and the small isorhizae. They were not
abundant, however, were difficult to analyse in pre¬
served material, and he had rather young animals.
Desmonemes seem to occur in the polyp stage only.

Records from New Zealand: SE of Banks Peninsula,
Christchurch (Kramp 1965); off Fiordland (Bouillon
1995, medusa); 300 km east of Cape Palliser (this
study); polyps not recorded yet.

Other Records (medusa): Mediterranean; Atlantic
Ocean from Bermuda to South Africa and Patagonia;
East Africa; Sri Lanka; Central North Pacific, Califor¬
nia; southern Japan; Vietnam; Philippines; Rarotonga;

east of Australia (Kramp 1968, map in Bleeker & van
der Spoel 1988).

Pandeopsis Kramp, 1959

Small, colonial hydroids with three filiform tentacles.
Medusa with the characteristics of the family, and
voluminous, quadrangular manubrium with large
base attached to subumbrella. Long mesenteries.
Gonads on interradial walls of stomach, smooth, sheet¬
like. Mouth with four simple lips. Up to 12 marginal
tentacles and up to 24 rudimentary bulbs. Tentacular
cirri or reduced tentacles absent. Tentacle bulbs
without spur, with abaxial ocelli.

Type Species: Pandeopsis ikarii (Uchida, 1927).

Fig. 42. Patidea conica . a) adult medusa after preserved material; scale bar 2 mm. b) nematocysts of medusa: basitrichous
isorhiza (?), microbasic eurytele, atrichous isorhiza; scale bar 10 |im. c) polyp stage, redrawn after Picard (1956), no scale
given; note: this polyp has not yet been found in New Zealand waters.

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Remarks: The life cycle of the only species of this genus
was revealed by Bouillon (1985b).

Pandeopsis ikarii (Uchida, 1927) (Fig. 43a-b)

Tinrnnnn ikarii Uchida, 1927: 208, fig. 35.

Pandeopsis scntigera Kramp, 1959: 232, fig. 7.

Pandeopsis scntigera Kramp 1961: 113.

Pandeopsis ikarii: Kramp 1961: 444; Kramp 1968: 41, fig. 105;
Bouillon 1985b: 257, fig, 6.

Material Examined:

No material from New Zealand seen.

Description:

Medusa stage (after Uchida 1927 and Kramp 1961):
Medusa up to 4.5 mm high, almost globular, jelly thick,
especially in apical region. Manubrium about half
height of bell cavity. Stomach short and very broad,
quadrangular, its entire surface attached to the sub¬
umbrella. The perradial edges of the stomach in their
entire length adnate to the radial canals, thus forming
mesenteries. Mouth rim almost smooth, with 4 simple

Fig. 43. Pandeopsis ikarii. a) medusa, modified after Uchida
(1927); scale bar 1 mm. b) polyp stage with stolon and
developing dispersal bud (right); redrawn from Bouillon
(1985b) with permission of the publisher; scale bar 0.2 mm.

lips. Four interradial gonads on stomach, each like a
flat sheet with trapezoid outlines, completely smooth.
The distal, free portions of the radial canals short and
narrow. Circular canal and velum narrow. Eight
marginal tentacles, each with an elongated pear-
shaped abaxial ocellus; no abaxial spur; with 8 small,
adradial rudimentary bulbs also bearing ocelli. In
living specimens each gonad is provided with 3 or 4
small red spots. Nematocysts (after Bouillon 1980):

a) reniform microbasic euryteles, (7-9) x (3.5-4.6) pm.

b) elliptic microbasic euryteles, (10) x (4.6) pm.

Polyp stage (after Bouillon 1985b, not known from
NZ): Planulae aggregate and attach to the substratum
with one end, thus forming a common hydrorhiza.
The other ends of the planulae each form a small polyp
(up to 0.4 mm). These with conical hypostome and
below it 1 whorl of 3 filiform tentacles. Polyps from
aggregates multiply by budding from the common
hydrorhiza. This hydrorhiza also forms long stolons
that form dispersal buds at their end. Single planulae
form solitary polyps only. Medusae buds unknown.

Type Locality: Seto, Japan.

Remarks: The description of Uchida (1927) and Kramp
(1961) slightly disagree on the surface structure of
the gonads. Uchida described them as complicated
and irregular. This discrepancy may eventually be
explained by the younger stages examined by Uchida.
Bouillon (1980) found this medusa in very large
numbers in Papua New Guinea.

Records from New Zealand: Kermadec Islands (Kramp
1965, medusa).

Other Records: Japan; Philippines; Gulf of Siam; Java
Sea; Papua New Guinea (Kramp 1968; Bouillon 1980).

Merga Hartlaub, 1913

Hydroids (where known) colonial, arising from
tubular ramifying hydrorhiza, cauli slightly branched
or not; with or without pseudohydrotheca which,
when present, not enveloping tentacles; one whorl of
filiform tentacles. Medusae buds arise from stems or
stolons. Pandeid medusae with perradial edges of
stomach connected with radial canals by mesenteries,
stomach with cross-shaped base, manubrium not
twisted, with simple or faintly crenulated oral lips.
Smooth adradial or interradial gonads. Four to eight
or more long tentacles, with or without rudimentary
bulbs or tentaculae, with or without ocelli (after
Bouillon 1980, 1985a).

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(Fig. 44ad)

Type Species: Merga violacea (Agassiz & Mayer, 1899).

Remarks: The difference between the diagnoses of
Merga and Pandeopsis is rather small and relies mostly
on the shape of the stomach.

Only one species is known from New Zealand.

Merga treubeli n.sp.

Material Examined:

2 juvenile medusae from Evans Bay, surface plankton, taken
12.1.1994 and 25.3.1994, cultivated to maturity.

5 juvenile specimens from surface plankton at Seatoun
(Wellington Harbour), 13.1.1994, cultivated to maturity
females and males obtained; gametes were spawned and
developed into planulae which did not metamorphose.
One female selected as holotype H-648, remaining material
deposited as paratypes, P-1076.

Fig. 44. Merga treubeli n. sp., from life, a) to c) all the same scale, also oral view of manubrium, a) youngest stage found in
the plankton, below oral view of manubrium; scale bar 1 mm. b) immature female from plankton, most frequently found
stage, c) mature male medusa cultivated from a similar stage as given in b). d) nematocysts: microbasic eurytele from
tentacles, same discharged, microbasic eurytele from mouth margin; scale bar 10 |jm.

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Description: Medusae with spherical umbrella, up to
4 mm high and wide, with very thick jelly; apical jelly
can reach half of the total height, dilated velum spans
two-thirds of radius. Manubrium half as long as bell
cavity, mouth small and cruciform. Base of manu¬
brium with 4 triangular, laterally compressed per-
radial extensions which continue as radial canals. At
beginning of each radial canal a triangular process
of variable height projecting upwardly into apical
mesogloea. Radial canals smooth. With 4 perradial, 4
interradial, and 8 shorter adradial tentacles; in adults
no rudiments of bulbs or tentaculae. Tentacles hollow,
evenly covered with nematocysts, bulbs small, without
ocelli. Gonads cover basal extensions of stomach in 8
adradial sheets which are in contact interradially but
are separated perradially. Nematocysts:

a) microbasic euryteles from tentacles, (6.5-7) x (3-
3,5) pm, s = 1.

b) microbasic euryteles from mouth, only few present,
(8.5-9) x (4.5-5) pm.

Size of spawned eggs: 102 pm (n = 5, one animal).
Colour: orange bulbs in nature.

Type Locality: Seatoun jetty, Wellington, New Zealand.

Remarks: With its small mouth and the triangular
projections at the proximal end of the radial canals M.
treubeli is quite distinct from all other Merga species.
The extensions of the stomach base were here inter¬
preted as mesenteries, although they are not typical.
Normally the stomach is more voluminous. It may be
necessary, therefore, to transfer this species to another
genus in the future, but further life-cycle information
is needed to substantiate this.

Etymology: The species name treubeli was chosen to
honour and thank the Treubel Foundation (University
of Basel) for financing this study.

Records from New Zealand: Wellington Harbour.

Bamettia n. gen.

Pandeid medusa with eight hollow, long tentacles
between each pair of which are cirri-like small ten¬
tacles without bulbs, with chordoid gastrodermis. The
cirri-like tentacles are evenly spaced and not associ¬
ated with the larger tentacles. Manubrium small, with
four simple perradial lips. Gonads interradial, smooth.
Four radial canals present, without mesenteries. Apical
projection may be present. Ocelli lacking. Cnidome
includes microbasic euryteles.

Type Species: Bamettia caprai n.sp.

Remarks: See below under Bamettia caprai, the only
species of this genus.

Bamettia caprai n. sp. (Fig. 45a-b)

Material Examined:

20 medusae collected by T. Barnett during March to July
1983 and 1984, One medusa selected as holotype and
deposited as H-639; some of the remaining material
deposited as paratype P-1067.

Description: Medusae up to 2 mm but mostly smaller,
bell shape variable from as broad as high to higher

Fig. 45. Bamettia caprai n.gen. n.sp., from preserved
material, a) mature medusa; scale bar 0.2 mm. b) nemato¬
cysts: larger microbasic heteroneme, microbasic eurytele;
scale bar 10 [im.

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than broad. Jelly thick, forming a blunt apical pro¬
jection. Manubrium about half length of bell cavity,
with 4 simple perradial lips without nematocysts.
Four large, interradial gonads covering almost the
whole manubrium. Gonads with a smooth surface and
margin. Four radial canals, moderately thin and
smooth. Ring canal broader than radial canals. No
mesenteries present, or these only indicated. In adults
4 perradial and 4 interradial long tentacles, length up
to 1 mm (preserved material). Tentacles with large
conical bulbs tapering into the tentacles which are
proximally hollow and distally filled with paren-
chymatic gastrodermis. Between each pair of these
large tentacles 2 (sometimes 3) evenly spaced, small,
cirri-like tentacles with chordoid gastrodermis. No
ocelli present. Nematocysts:

a) larger microbasic heteronemes (mastigophores or
euryteles), (11.5-13.5) x (4-5) j.im.

b) smaller microbasic euryteles, (6.5-7.5) x (2-3) pm,
s = 1.

Polyp stage: unknown.

Type Locality: Surface plankton near Goat Island,
Leigh Marine Reserve, New Zealand.

Remarks: With its cirri-like tentacles and smooth
gonads, the morphology of Barnettin cnprai n.gen. n.sp.
superficially resembles medusae of the genus Hnlitinrn.
Barnettin cnprai always has more than 4 large ten¬
tacles, although in juveniles the interradial ones are
present as bulbs only. Hnlitinrn belongs to the family
Protiaridae (see p. 76 and Bouillon 1985a), character¬
ised by having only 4 normal perradial tentacles.
Additionally, the examined species of this family have
a cnidome and polyps that are quite different from
the ones found in the Pandeidae (Bouillon 1980,1985a,
1988). It appears that the Protiaridae are not closely
related to the Pandeidae. Because B. cnprai has more
than 4 normal tentacles and microbasic euryteles, the
genus is included in the Pandeidae. In this family the
only genus that has cirri-like tentacles is Cirrhitinrn
Hartlaub, 1913, which is, however, otherwise well dis¬
tinguished by its folded gonad, mesenteries, and ocelli.
A new genus is thus required.

Barnettin cnprai seems to be a rather abundant
medusa near Leigh as Barnett (1985) collected about
130 animals. Barnett's data indicate that they are
seasonal as the medusae were found in late summer
to autumn only (February to July).

The life cycle of this species has yet to be revealed.
Information on the polyp stage will also be needed to
confirm its affinities with the family Pandeidae.

Etymology: The genus name was chosen to honour and
thank Mrs Treffery Barnett for the generous loan of

her collection of Anthomedusae and allowing the neu
species to be described here. The species name cnprai
refers to the type locality of Goat Island (cnprn, Latin
goat).

Records from New Zealand: Goat Island (Leigh)
Whangateau Harbour (Barnett 1985, as Hnliterelln sp.)

Family PROTIARIDAE Haeckel, 1879

Medusa with only four fully developed tentacles
arising from well-developed hollow tentacular bulbs
Manubrium with four simple lips. Four interradial
gonads with smooth surface. With or without mesen¬
teries. With four simple radial canals and a circular
canal. Margin may have small cirri-like tentacles
exceptionally with ocelli (after Bouillon 1985a).

Remarks: Only Hnlitinrn is known from New Zealand
Halitiara Fewkes, 1882

Protiarid medusae with four simple perradial tentacles
and in between them numerous solid cirri-like
tentacles. With or without mesenteries, with or without
apical projection. Lacking ocelli (after Bouillon 1985a)
Type Species: Hnlitinrn formosa Fewkes, 1882.

Remarks: The life cycle of only H. inflexn is known
(Bouillon 1985b). Two species of Hnlitinrn have been
recorded from New Zealand. Characteristics are:

H. formosa: with apical projection, without mesen¬
teries.

H. inflexn : lacking apical projection, mesenteries
present.

Halitiara formosa Fewkes, 1882

Halitiara formosa Fewkes 1882: 267, pi. 4, fig. 2.

Protiarn formosa : Mayer 1910: 107, pi. 6, figs 4-6.

Halitiara formosa : Kramp 1959: 115, fig. 103; Kramp 1961:
102; Kramp 1965: 28; Kramp 1968: 40, fig. 102
Brinckmann-Voss 1970: pi. 11, fig. 1; Goy 1972: 982, figs
5, 7; Bouillon 1995: 230, fig. 5.

Material Examined:

Several medusae from NZOI Stns N401 and N413, identified
by Prof. J. Bouillon as including H. formosa (see Bouillon
1995).

Description (after Kramp 1968, Bouillon 1995): Medusa
up to 3 mm high, pear-shaped, with solid apical pro¬
jection; manubrium pyriform, about half as long as

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bell cavity; 4 long, hollow and 24-35 short, solid
tentacles, tightly coiled, cirrus-like; no ocelli; gonads
interradial, smooth; no mesenteries.

Type Locality: Tortugas, Florida.

Remarks: Kramp (1965) was the first to record Halitiara
formosa from New Zealand. He found only one
medusa, that did not entirely accord with other popu¬
lations owing to its crenulated lips. He therefore
considered this identification as doubtful. Kramp's
sample is no longer available. Bouillon (1995) identi¬
fied several medusae from New Zealand as H. formosa
and illustrated one of them. His samples were re¬
examined for this study and I conclude that they do
not contain H. formosa. Some of the medusae resemble

H. formosa in general appearance, including the cirri¬
like tentacles, but their gonads arc in obvious folds
which are directed perradially. A closer examination
additionally revealed remainders of nematocyst tracks
on the exumbrella. The medusae thus resemble some
Leuckartiarn species. The samples are not in good con¬
dition and a more precise identification was not
possible. The presence of H. formosa in New Zealand
cannot therefore be confirmed and all records must
be considered uncertain. Judging from the distribution
of the species, however, it may be expected to occur
in New Zealand waters.

Records from New Zealand: ? some distance off the
east coast of South Island (Kramp 1965); ? off Foxton,
Cook Strait; ? south of Kaikoura (Bouillon 1995).

Other Records: Tortugas, Florida; Bahamas; Medi¬
terranean; Japan; India; Papua New Guinea; ? Fiji
(Kramp 1961, Goy et al 1972, Bouillon 1980).

Halitiara inflexa Bouillon, 1980 (Fig. 46a-b)

Halitiara inflexa Bouillon 1980: 324, fig. 9; Bouillon 1985b:
259, fig. 7; Bouillon et al. 1988b: 212, fig. 8; Goy et al.
1991: 110, fig. 27; Bouillon 1995: 230, fig. 6.

Material Examined:

4 medusae from NZOl Stn N404, 41 0 38 f S, 175°18.8'E, (Cape
Palliser), 17.12.1974, 0-51 m.

Description:

Medusa stage: Umbrella bell-shaped, 1.6 mm high and

I. 2 mm in diameter. Jelly moderately thick, gradually
thickening towards top to about twice the thickness
of the lateral walls. Manubrium voluminous, quad¬
rangular, length about two-thirds of bell cavity, joined
to radial canals by mesenteries for half of their length.
Mouth with 4 simple lips. Gonads large, bulging, fill-

Fig. 46. Halitiara inflexa. a) medusa from Stn N404; scale
bar 0.25 mm. b) polyp stage, redrawn from Bouillon (1985b)
with permission of the publisher. Note: the polyp stage has
not yet been found in New Zealand. Scale bar 1 mm.

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ing interradial position completely, leaving free only
a small perradial band of stomach and the mouth
region. Four radial canals and circular canal, all narrow
and with smooth margins. Four long perradial ten¬
tacles. Tentacles with broad conical base then tapering,
base not laterally compressed. Nematocysts evenly
distributed on tentacles. Between each pair of long
tentacles 3-6 short, cirri-like tentacles, without bulbs,
often coiled. Cirri with chordoid endodermis, tips with
nematocysts (haplonemes). Nematocysts (after
Bouillon 1980, Bouillon et al 1988b):

a) larger atrichous isorhizas, (10) x (4.5) pm.

b) smaller atrichous isorhizas, (7) x (2) urn.

c) merotrichous isorhizas, (15) x (9) pm.

d) mastigophores, (7) x (2.5) pm.

Polyp stage (after Bouillon 1985b, not known from
NZ): Hydroid colonies arising from attached, ramified
stolons. Stolons covered by perisarc. Polyps up to
1 mm, with a very short caulus (80 mm), a long, nar¬
row cylindrical body and a short conical hypostome.
Below hypostome one whorl of 10 long, filiform ten¬
tacles (0.7 mm) with irregular clusters of nematocysts.
Alternating with the tentacles are large nematocysts.
Caulus and base of polyp body are covered by a
perisarc cup (height 0.17 mm) into which the polyp
can almost completely retract.

Type Locality: Laing Island, Papua New Guinea.

Remarks: Halitiara inflexci from New Zealand agrees
well with the original description by Bouillon (1980),
only the size is somewhat smaller. The nematocysts
were examined for this study but, owing to poor pre¬
servation, it was not possible to recognise them clearly.
It was evident however, that the cirri contain a
concentration of larger haplonemes at their tips.

Records from New Zealand: Cape Palliser (Bouillon
1995).

Other Records: Papua New Guinea, Mediterranean
(Bouillon 1980; Goy et al. 1991).

Family EUDENDRIIDAE L. Agassiz, 1862

Colonial hydroids with an erect, usually branched
stem enclosed by firm perisarc up to the base of the
hydranth body. Hydranths large, with trumpet¬
shaped hypostome and one or more whorls of filiform
tentacles immediately below it. Reproduction by fixed
sporosacs borne on the hydranth body below the
tentacles, reproductive hydranth often reduced to a
blastostyle. Male gonophores usually with several

chambers in linear series. Young female gonophore
with a single egg encircled by a spadix (after Millard
1975).

Remarks: Members of this family are easily dis¬
tinguished from all other families of the Athecata by
their wide, trumpet-shaped hypostome. The
hydranths in this family are comparatively large and
urn-shaped. The perisarc originates in a circular grove
with larger cells at the base of the hydranths (basal
furrow). Initially the perisarc is very thin, becoming
thicker only on the cauli (Fig. 48b). The gonophores of
this family show no trace of a medusa stage (Wasser-
thal 1973).

The family comprises the genera Myrionenm and
Eudendriuni, which differ in the number of tentacle
whorls. Only the genus Eudendriwn is known in New
Zealand.

Eudendriwn Ehrenberg, 1834

Colonial hydroids with monopodial growth, hydro-
caulus enclosed by perisarc. Hydranths on perisarc-
covered cauli, radially symmetrical with large, under¬
cut hypostome and only one whorl of filiform tentacles.
Microbasic euryteles always present in tentacles, other
kinds of nematocysts may be present on hydranth
body, hypostome, or gonophores. Gonophores fixed
sporosacs, borne on hydranth body below tentacles.
Reproductive hydranth often reduced to a blastostyle.
Male gonophores with one or more chambers in linear
series. Young female gonophores with a single egg
encircled by a spadix.

Type Species: Eudendriwn rnniosuni (Linnaeus, 1758).

Remarks: Eudendriwn is easy to recognise, but identi¬
fication to species can be difficult. Earlier authors
often based species distinctions on traits that are now
known to be too variable or dependent on the environ¬
ment (see Millard 1975). The use of nematocyst
characters has improved the situation.

Most Eudendriuni species have limited larval
dispersal (Sommer 1992). Possibly speciation rates
have been high, for many endemic species exist (cf
Watson 1985; Hirohito 1988; Marinopoulos 1992).

The New Zealand Eudendriuni species were
recently revised by Watson (1987). Watson (1985)
reviewed the Australian Eudendriuni species some of
which may yet occur in New Zealand. For the identifi¬
cation of Eudendriwn species nematocysts must always
be examined. The four known species of Eudendriun:
occurring in New Zealand can be distinguished with
the help of nematocysts only, and more species may

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be expected to occur. Characteristics of known New

Zealand species:

Eiidendrium novaezelandiae: normally large colonies,
fascicled, two sizes of nematocysts in hydranth, the
larger eurytele more than twice as large as the
smaller, its shaft not longer than capsule.

Eiidendrium ritcliiei: completely annulated perisarc, two
types of euryteles, the larger with everted shaft
much longer than capsule, female gonophore with
bifurcated spadix.

Eiidendrium terranovae : large, fascicled colonies, only

one size of euryteles.

Eiidendrium maorianus : small colonies, only sporad¬
ically annulated, two sizes of euryteles, the larger cap¬
sules smaller than twice the size of the smaller cap¬
sules, shaft of larger capsules not longer than capsule,
spadix of female gonophore 0-2 times branched.

Eiidendrium novaezelandiae Marktanner-

Turneretscher, 1890 (Fig. 47a-f)

Eiidendrium novnezelnndine Marktanner-Turneretscher 1890:
201, pi. 3, fig. 21; Watson 1987: 326, figs 14.

Fig 47. Eudendrium novaezelandiae , from life except e) and f). a) distal part of colony from Kaikoura; scale bar 2 mm. b)
hydranth from Mahanga Bay, Wellington Harbour; scale bar 0.5 mm. c) nematocysts: small microbasic eurytele from tentacles,
same discharged, large microbasic eurytele from body and hypostome, same discharged; scale bar 10 pm d) young female
gonophore from Te Raekaihau; scale bar 0.2 mm e) mature female gonophores from Stn C380, same scale as d). f) male
gonophores from Stn A439, same scale as d).

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Material Examined:

1 colony from NZOI Stn A439, 40°59.6’S, 174°27.2E, Cook
Strait, 4.10.1958, 140-160 m, small colony on polychaete
tube, with male gonophores.

2 colonies from NZOI Stn C380, 38°50'S, 174 0 21.5'E, north
of New Plymouth, 37 m, 28.10.1959, 25 and 40 mm high,
hydranths mostly lost, strongly fascicled, both fertile
females.

1 microslide prepared by P. Ralph with terminal portion of
colony, labelled: Eudendrium spec, from Ikntere, drift,
26.5.1950. Locality unknown; colony bears female gono¬
phores with various degrees of hydranth reduction.

1 live colony from Kaikoura, dredged from 90 m, March
1994, few hydranths and no gonophores left, 60 mm high,
cultivated in running sea water for 1 month then pre¬
served, deposited.

2 very small, live colonies with few hydranths, collected
13.3.1994 east of Te Raekaihau, 2 m, on sponges, one with
developing female gonophores.

1 very small Jive colony with only 2 hydranths, infertile,
from Mahanga Bay, Wellington Harbour, 27.1.1994, 1 m.

Description: Hydroid colonies with erect branching
stems arising from attached ramifying stolons. Colony
may reach 60 mm in height. Larger colonies fascicled
almost to the tips, branching irregularly in various
planes. Perisarc of erect parts with dispersed intervals
of annulation, otherwise smooth. Hydranths with 20-
24 tentacles, top of hypostome often rather flat in life,
without cnidophores but with conspicuous nemato-
cysts on hypostome and as a ring on basal part of
hydranth body, Male and females on separate colonies.
Male gonophores develop on blastostyle without ten¬
tacles, in groups of up to 12, each with 1-2 chambers,
some with a distal knob containing nematocysts.
Female gonophores develop initially from a smaller
hydranth with reduced tentacle number. Spadix of
female gonophores initially present, simple, not
branching. Tentacles and spadices are lost during later
development. Nematocysts:

a) large microbasic euryteles, on body, hypostome
and gonophores, discharging at 45° angle to main
axis, with spines, (18-31) x (8-15) mm, s = 0.8-0.9.

b) intermediate microbasic euryteles, rare, present in
female gonophores only, (11-17) x (4.5-8.5) mm,
s = ~ 0.9.

c) small microbasic euryteles, on tentacles, dis¬
charging almost directly foreward, (6.5-9.5) x (2.5-
4) mm, s = 0.8-0.9.

Colours: live hydranths have a characteristic creamy
to yellow colour; stems brown, becoming lighter
distally.

Type Locality: New Zealand.

Remarks: The identification of small infertile E. novae-
zelandine colonies is based mainly on the presence of

the conspicuous large microbasic euryteles in addition
to the smaller ones in the tentacles. The size ratio
between these capsules is always larger than 2.5
although the large capsule especially has a very vari¬
able size in different colonies. The capsule types and
sizes observed in this study are as described by Watson
(1987), but their size range was wider.

The colony size of fertile animals varied con¬
siderably, from only a few hydranths to several dozen
The small colonies, mostly infertile, were obtained
from shallow depths, while all larger ones were from
deeper water (37-140 m).

The colour of the live hydranth was found to be
very characteristic and allowed a quick preliminary
identification. This specific colour was even observed
to some extent in alcohol-preserved animals and in
cultivated animals fed on Artemi a.

Records from New Zealand: Coromandel (Watson
1987), Taranaki Bight, Cook Strait, Wellington, Kai¬
koura (this study).

Ot her Records: Not known outside New Zealand.

Eudendrium ritchiei Millard, 1975 (Fig. 48a-f

Eitdendriuni insigne : Ralph 1953: 63, fig. 2A-B.

[Not Eudendrium insigne Hincks, 1861]

Eudendrium ritchiei Millard 1975: 87, fig. 30; Watson 1987
327, figs 5-8.

Material Examined:

Live colonies from Cheltenham Beach, Auckland, on drift
seaweed, 27.7.1991, infertile.

Live colonies from Narrow Neck Beach, Auckland, on Cnrpe-
phylluni maschalocnrpuni, 1 m, 7.1.1994, male and female
colonies.

Description: Hydroid colonies with erect branching
stems arising from loosely adhering, ramified, thick
stolons. Stolons not annulated. Stems reaching 15 mm.
in height, not fascicled in New Zealand specimens
branching irregularly in all planes but with a tendency
to unilateral growth. Perisarc of erect parts strongly
annulated throughout. Hydranths with 17-25 tentacles
and no cnidophores, but a ring of large euryteles
towards base of hydranth above basal furrow. Males
and females on separate colonies. Male blastostyles
with no tentacles, bearing a cluster of one-chambered
gonophores with an occasional terminal tubercle
containing large euryteles. Hydranths developing
female gonophores with reduced tentacles which may
be lost in later development, 3-6 gonophores in a
whorl. Young female gonophores with bifurcating
spadix clasping the single egg, older gonophores oval

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with a smooth transparent capsule and without spadix,
Nematocysts and other measurements:

a) larger macrobasic euryteles, present as ring on
body of hydranth and on male and female gono¬
phores, shaft with spiral bands of spines, (17.5-
21.5) x (6.5-9) ,11m, s = 2.8.

b) smaller microbasic euryteles, present in tentacles,
(6.5-7.5) x (3.5-3.5) pm, s = 0.8.

Caulus diameter 110-140 pm, stolon diameter 155-
190 pm. Colour of hydranths white to grey, sometimes

slightly green, gonophores yellow.

Type Locality: Saunders Rocks, Sea Point, South Africa.

Remarks: The material of E. ritchiei from New Zealand
available for the present study agrees in most respects
with the diagnostic features given by Millard (1975).
The main difference, as in the material examined by
Watson (1987), is that the New Zealand colonies are
not fascicled like the South African ones. This may
depend on the environment and age however.

Fig. 48. Eitdendriitni ritchiei from preserved material, a) colony form, note strong annulation of perisarc, CL; scale bar 1 mm.
b) hydranth showing belt of large euryteles (arrow) and below the basal furrow, CL; scale bar 0.2 mm. c) nematocysts:
macro- and microbasic euryteles; scale bar 10 jjm. d) young female gonophore showing characteristic bifid spadices (shaded),
CL, same scale as b). e) older female gonophores with lost spadices, same scale as b). f) male gonophores, the arrow points
towards distal knob with euryteles, same scale as b).

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Records from New Zealand; East coast beaches from
Auckland to Gisborne (Ralph 1953, as E. insigne), Coro¬
mandel (Watson 1987), Hauraki Gulf (this study).

Other Records: South Africa (Millard 1975).

Eudendrium terranovae Watson, 1985 (Fig. 49a-d)

Eudendrium sp.: Totton 1930: 141.

Eudendrium terranovae Watson 1985: 189, figs 20-23.

Material Examined;

2 colonies from NZOI Stn E413, 45°12'S, 171°44'E, near
Oamaru, 11.10.1965, 594 m (error ?), Agassiz trawl; 4 and
10 cm high colonies, one colony a fertile male, det. by J.
Watson.

1 male colony from Cliffy Island, Australia, MV no. F50503,
cited in Watson (1985).

1 female colony from Cliffy Island, Australia, MV no.
F50504, cited in Watson (1985).

Description: Hydroid colonies with erect branching
stems to 10 cm height, arising from attached, ramified
stolons. Main stems fascicled. Branching rather
regular, mostly in one plane. Perisarc at origin of new
branches may have 0-9 annulations, otherwise smooth.
Hydranth with 22-28 tentacles. Male and female gono-

phores on separate colonies. Mature male gonophores
small, on a blastostyle devoid of tentacles at all stages,
up to 30 in tightly-packed clusters at the end of a
caulus, with up to 3 chambers, immature gonophores
with terminal knob. Female gonophores borne on a
blastostyle showing atrophy of the tentacles from early
growth stages, tentacles completely absent at maturity.
Spadices in younger female gonophores present, not
branching. Nematocysts:

a) microbasic euryteles, discharging directly forward,
with indistinct terminal swelling of shaft, (6-
9) x (2.5*4) |Lim, s ~ 0.8.

Measurements: stolon diameter 160-240 pm; diameter
stems at base of colony 220-250 pm; diameter of cauli
140-200 pm.

Type Locality: North Cape, New Zealand.

Remarks: The preferred branching in one plane is very
characteristic of E. terranovae.

Records from New Zealand: North Cape (Totton 1930),
Oamaru (this study).

Other Records: Cliffy Island (Australia).

Fig. 49. Eudendrium terranovae f from material from Cliffy Island (Australia) except b). a) whole colony, sample F50503,
scale bar 5 mm. b) nematocysts: microbasic eurytele, same discharged; scale bar 10 pm. c) female gonophores from sample
F50504; scale bar 0.2 mm. d) male gonophores from sample F50503, same scale as c).

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Eudendrium maoriamis n.sp.

(Fig. oOa-f)

Material Examined:

Small colony from western side of Te Raekaihau (Welling¬
ton), 1 m, 13.1.1994, erect parts once or twice branched,
male gonophores developed in culture.

1 colony from western side of Te Raekaihau, 1 m, 30.1.1994,
on sponge, 3 times branched, infertile.

Several colonies from western side of Te Raekaihau, 2 m.

26.2.1994, mature female colony on stone, 10 mm high,
deposited as holotype H-642. One male colony, stem ~
10 mm, 3-4 times branched, deposited as paratype P-1083.

Several colonies from South Bay, Kaikoura, South Island,
1 m, 1.3.1994, one female colony, 10 mm high; one male
colony, 10 mm high, 2-4 terminal hydranths, material
deposited as paratype P-1069.

1 colony from western side of Te Raekaihau, intertidal,
13.3.1994, 18 mm high, infertile.

Fig. 50. Eudendrium maoriamis n.sp, from life, a) type specimen, female colony; scale bar 1 mm. b) hydranth; scale bar
0.5 mm. c) nematocysts: microbasic euryteles; scale bar 10 urn, d) male gonophores; scale bar 0.2 mm. e) mature female
gonophores, the hydranth and spadices are completely lost and the caulus has resumed growth distally, same scale as in d).
f) younger female gonophores that show the variation of the spadix form (shaded), same scale as d).

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cACt

1 colony from western side of Te Raekaihau, 2 m, female
colony on sponge, together with Hybocodon prolifer .
Numerous colonies from underneath Queens Wharf, Well¬
ington Harbour, subtidal, 16.6.1994, very abundant on
oysters, some fertile.

Description: Hydroid colonies with erect branching
stems arising from attached, ramified stolons. Stems
reaching 18 mm in height, branching sparingly, with
up to 10 terminal hydranths per erect stem, mostly
only 3. Perisarc of erect parts annulated at intervals.
Hydranths with up to 16 tentacles. Male and female
gonophores on separate colonies. Male blastostyles
with no tentacles at any stage of development, bearing
a cluster of gonophores with 1-3 chambers and an
occasional terminal tubercle containing euryteles.
Female gonophores initially on small hydranth with
tentacles which is later completely reduced, up to 6
gonophores per hydranth, spadix long and coiled,
often bifid, but in the same colony they may also be
unbranched or twice branched. Spadices are lost in
later stages and the egg is covered by a thin perisarc
hull. In mature blastostyles the caulus may resume
growth distally. As nematocysts there are two distinct
types of euryteles with an observed size ratio of 1.28-
1.46 (mean = 1.35, s.d. = 0.075, n = 7 colonies):

a) larger microbasic euryteles on body and male gono¬
phores, shaft with distinct spines, (9-12) x (3-5) pm,
s = 0.8.

b) smaller microbasic euryteles on tentacles, shaft with
delicate spines, (6.5-9) x (2.5-4) pm, s = 0.8-0.9.

Colour of living hydranths: orange to pink.

Type Locality: West side of Te Raekaihau, Wellington.

Etymology: The species name refers to the first inhabi¬
tants of New Zealand, the bifid and coiled spadix being
somewhat reminiscent of Maori symbolic motifs (Fig.
50f).

Remarks: Eudendriiun maorianus seems to be quite
abundant and widespread. It would be interesting to
determine whether it occurs only in the more southern
parts of New Zealand, replacing E. ritchiei which seems
to be confined to more northerly regions.

The low stems with few hydranths, the two
different euryteles, and the long and occasionally
branched spadix are the main distinctive characters
that separate £. maorianus from the previously known
New Zealand species. Although the total size range
of the two capsules observed in several populations
overlaps slightly, the size ratio between the two
capsule types within one colony was always larger
than 1.28 and both types are easily distinguishable,
even without measurements.

Records for New Zealand: Wellington, Kaikoura.

Family PROBOSCIDACTYLIDAE

Hand & Hendrickson, 1950

Colonial hydroids connected by stolons which are not
covered by perisarc. Hydranths sessile and poly¬
morphic: with gastrozooids and gonozooids, some¬
times dactylozooids. Gastrozooids with only two
filiform tentacles originating below a large hypostome
which is densely covered by nematocysts. Gonozooids
and dactylozooids without mouth and tentacles.
Gonozooids produce free medusae. Medusae with a
manubrium that has four, six, or more gastric lobes
that extend along the proximal parts of the radial
canals. Radial canals may be branching and an
obliterated radial canal may be present or completely
absent. The gonads encircle the manubrium and can
extend onto the gastric lobes. Alternating with the
tentacles may be clusters of nematocysts on the
exumbrella. Ocelli and statocysts absent.

Remarks: This diagnosis was amended after Bouillon
(1985a) to allow the inclusion of the new genus
Fabienna. The emendation concerns the possible
absence of exumbrellar nematocyst clusters. The
Proboscidactylidae have been included in the Limno-
medusae by some authors (e.g., Kramp 1961; Bouillon
1985a). The absence of statocysts, the polyps with
stolons, and the presence of desmonemes indicate that
this family is better placed in the Filifera of the Antho-
athecata. Other authors have already proposed this
classification (Werner 1984; Petersen 1990). The
inclusion of Fabienna is somewhat of a compromise
and the arguments for it are discussed below.

Diagnostic characters for genera known from New
Zealand:

Fabienna : medusa without medusa buds on manu¬
brium, without exumbrellar nematocyst pads,
nematocysts of tentacles confined to tip.
Proboscidactyla : medusa with medusa buds on manu¬
brium, with patches of nematocysts on exumbrella,
tentacles covered with nematocysts along entire
length.

Fabienna n.gen.

Medusae with slightly lobed umbrella margin. With
four perradial tentacles that have their origin some¬
what displaced towards the exumbrella. Larger
nematocysts confined to tentacle tips in one terminal
cluster immediately followed proximally by an adaxial
cluster; the two clusters may fuse in older individuals.
Cnidome includes macrobasic euryteles. Gonads
develop on manubrium only, in an interradial position.
Circular canal present as solid cord.

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Type Species: Fnbiennn s pi meric a n.sp.

Remarks: The new genus is provisionally included in
the Proboscidactylidae. Possible synapomorphies are
the solid radial canal and the macrobasic euryteles.
Influencing this decision was the inclusion of
Pochella oligonemn Kramp, 1955 in this family. Pochella
oligonemn is congeneric with Fnbiennn.

Etymology: The genus name Fnbiennn is dedicated to
my little daughter Fabienne, in order to acknowledge
this source of much joy and inspiration.

Fabienna sphaerica n.sp. (Fig. 51a-g)

Material Examined:

2 juvenile specimens from Evans Bay, 14,7.1994, cultivated
until gonad tissue became visible (2.7.94).

1 juvenile specimen from Seatoun.

1 advanced but not mature specimen from Evans Bay,
30.8.1994, 1 ciliary field damaged and missing, preserved
after one day, deposited as holotype H-644.

1 juvenile specimen from Evans Bay, 1.9.1994, cultivated
until female gonads became visible (20.9.94).

1 juvenile and one fully mature animal, both damaged,
collected by T. Barnett near Goat Island, Leigh Marine
Reserve, date not known (1984?), not mentioned in her
thesis (1985), both medusae deposited as paratype P-1070.
For comparison, holotype and 2 paratypes of Pochella
oligonema Kramp, 1955, held by ZMC, collected 26.1.1946,
Atlantide Stn 77, Accra, Shana.

Also ~15 medusae of Kantiella euignmtica Bouillon, 1978a
originating from Laing island, Papua New Guinea,
collected and identified by Prof. J. Bouillon.

Description:

Mature medusa: Up to 1.8 mm high, bell rather
spherical. Jelly thick, apex about 1.5 times thicker than
lateral walls. Bell-margin lobed through 4 perradial
furrows where tentacles originate. Velum when dilated
spanning half of radius. On umbrella margin in each
interradial position a small triangular field with long
cilia. These fields are slightly more opaque than the
epidermis of the exumbrella. The base of these fields
lies along the circular strand. A few scattered nemato-
cysts are situated along the bell margin, occasionally
more near the base of the ciliated fields. Manubrium
about two-thirds as long as the bell cavity, with large
cruciform base, without a peduncle. Distal part of
manubrium simple, conical, mouth opening circular
to quadrangular, margin provided with nematocysts.
Gonads in the form of 4 large, interradial, triangular
pads leaving manubrium free only at small perradial
zone and near mouth. With 4 narrow, hollow radial
canals. Very fine, blindly ending lateral branches
rarely observed. Circular gastrodermal strand running

closely along bell-margin, thin and no lumen visible.
Four perradial marginal bulbs, egg-shaped, gastro-
dermis very opaque and probably without lumen,
below gastrodermis a pad of epidermis with nemato¬
cysts. The circular strand is in direct connection with
the gastrodermis of the bulb. Four tentacles arise some¬
what towards the exumbrella and may be adnate to it
for a short distance. Tentacles without terminal swell¬
ing, but terminal region often bent like a hook. Tentacle
gastrodermis proximally parenchymatous for a short
distance, without lumen, then chordoid. Tentacles
mostly without nematocysts, these concentrated at tips
of tentacles in one terminal cluster followed by another
cluster confined to the adaxial side. No ocelli or
statocysts present. Nematocysts (some also present in
subumbrella along radial canals and along circular
strand):

a) macrobasic euryteles, from tentacles, with spines
only terminally or all along the shaft (14.5-17) x (9-
10) pm, s = 6-9.

b) macrobasic euryteles from mouth, (12-17) x (6-
8) pm, s = 4.7.

c) small capsules in tentacle bulbs, rare, ev. isorhiza,
(5-6) x (3-3.5) pm.

d) microbasic euryteles, (8-11) x (3.5-5) um, s = 2.2.

Young medusa: - 0.9 mm high, spherical, with thick
jelly, jelly at apex almost twice as thick, indented as
funnel at apex, margin slightly lobed. Manubrium a
simple inverted cone, basal perradial extension not
present. Tentacles short and stiff, bent upwards,
without terminal swelling. Few nematocysts in only
one cluster at tentacle tip. Interradial ciliary field only
in some animals visible.

Type Locality: Greta Point, Evans Bay, Wellington.

Etymology: The species name refers to the almost
spherical shape of this medusa.

Remarks on Taxonomy: Fnbiennn splmericn can be placed
only with difficulty into the present taxonomic system.
The main reasons for its inclusion in the Proboscidac¬
tylidae are given above. The absence of exumbrellar
pads of nematocysts and the absence of desmonemes
may argue against this, but in other families such
variation can also be seen. The closest resemblance of
Fnbiennn splmericn to any known medusa is to Pochelln
oligonemn Kramp, 1955. Two species of Pochelln have
been described: P. polynemn Flartlaub, 1917 and P.
oligonemn Kramp, 1955. Edwards (1973a) was able to
determine the life cycle of P. pohjnemn and showed
that this medusa belongs to a hydroid known as
Tricky dm pudicn (see also Rees 1941), a senior synonym.
Based only on its superficial similarity and pending

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Fig. 51. Fabienna sphaerica n.gen., n.sp., from life except c), e) and f). a) very young medusa; scale bar 0.2 mm b) medusa,
just before maturation of gonads; scale bar 0.5 mm. c) manubrium with mature gonads from a preserved medusa from
Leigh, same scale as a), d) nematocysts: discharged macrobasic eurytele, same not discharged, small capsule, microbasic
eurytele, same discharged; scale bar 10 |im. e) tentacle tip of preserved holotype showing the characteristic bending, note the
terminal nematocyst cluster (arrow) and the adjacent adaxial cluster (arrowhead); scale bar 0.1 mm. f) frontal view of
tentacle bulb from preserved holotype, CL, fixation tends to accentuate the lobes of the bell margin; scale bar 0.1 mm. g)
lateral view of tentacle bulb of living medusa, CL, same scale as f). Abbreviations: bm bell margin, cs circular strand, ex
exumbrella, up nematocyst pad, rc radial canal, tb tentacle bulb, te tentacle.

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more information on its life cycle, P. oligonemn was
provisionally also transferred to Triclnjdrn by Edwards
(1973a). The type specimens of P. oligonemn Kramp,
1955 were re-examined for the present study. This
medusa is different from that of Trichydrn and must
be placed in a separate genus. It has 4 tentacles with
nematocysts present only in a terminal cluster and an
adjacent adaxial cluster, exactly as described for
Fnbiennn sphaericn (Fig. 51e). In older medusae they may
fuse. This arrangement was indicated by Kramp (1955)
in his figure 7. The medusa of 7. piulicn has many
tentacles covered throughout with nematocysts
(Edwards 1973a; unfortunately, the kinds of nemato¬
cysts of Trichydrn are unknown). In one of the para-
types of P. oligonemn, is an interradial triangular field
like the ciliary fields of F. sphaericn . Kramp's P.
oligonemn resembles Fnbiennn sphaericn so closely that
they arc certainly congeneric. Some small differences
between these two samples can be found. Kramp's
medusa has a peduncle (only half the size as given in
his figure of 1955). The tentacle bulbs are slightly
different; as in Kramp's medusae they become
narrower towards the circular strand. The bulbs are
therefore almost as in Knntielln (cf. Bouillon 1978a, c).
The gonads in Kramp's medusae are also oval in shape
compared to triangular in F. sphaericn. The prismatic
shape of the stomach in the holotype of P. oligonemn
may be a fixation artefact only, as the mouth is widely
gaping. In the paratypes the stomach is obviously
cruciform in section and the mouth has 4 very incon¬
spicuous lips provided with nematocyst concen¬
trations.

Because the type species of Pochelln has been trans¬
ferred to Triclnjdra, the former name cannot be used
anymore and Kramp's medusa is here included in the
new genus Fnbiennn as 7. oligonemn n. comb. More data,
especially life-cycle information, of both Fnbiemw
species are urgently needed to confirm their syste¬
matic position. The relationship to the Laingiomedusae
Bouillon, 1978c should also be elucidated. An exam¬
ination of Knntielln enigmnticn Bouillon, 1978a revealed
that younger individuals of this species have tentacle
tips that are astonishingly like the special tentacle tips
of Fnbiennn: one terminal cluster followed by an adaxial
one, with a bend after after the proximal cluster. The
two clusters are more widely separated than in Fnbi¬
ennn and in older individuals the nematocyst clusters
are mostly grown together, otherwise Knntielln and
Fnbiennn are distinct. Some of the major distinctions
in Knntielln are: the exumbrellar nematocyst bands, the
pronounced separation of the tentacular bulbs from
the circular strand, and the origin of the tentacles.

General Remarks: Few specimens of Fnbiennn sphaericn
were found in Evans Bay during the winter months.
The occurrence of very young stages in Wellington

Harbour may indicate that the polyp must be present
in the proximity of Wellington. Medusae are easy to
keep at room temperature and they feed vigorously
on Artemi a. Gonad tissues become recognisable quite
soon and the morphology does not change much.
Developing gonads in the cultivated medusae were
as in the mature medusa obtained from Leigh.

The interradial triangular fields on the umbrella
margin are known only from this species and are
clearly visible only in live specimens. These fields are
composed of epidermal cells with long cilia. Their
function remains unknown. Normally these fields do
not contain nematocysts, but some few may occur at
their base along the bell margin. The bell margin itself
normally has some scattered nematocysts. In the
preserved samples from Leigh with a rather damaged
and crumpled umbrella, the ciliated fields could not
be seen.

Because F. sphaericn contains comparatively few
nematocysts and only few specimens were available,
not enough nematocysts could be analysed. It seems
that all capsules are very variable, even within one
animal, and the various euryteles distinguished above
may actually be only extremes of a continuous row.

Records from New Zealand: Wellington Harbour;
Leigh Marine Reserve (this study).

Proboscidactyla Brandt, 1835

Colonial hydroids conforming to the family diagnosis,
living as obligatory commensals with polychaetes.
Medusae with a manubrium that has four, six, or more
gastric lobes extending along the proximal parts of
the radial canals. Radial canals often branching and a
solid circular strand may be present or completely
reduced. Gonads encircle the manubrium and extend
onto the gastric lobes. Alternating with the tentacles
the exumbrella bears clusters of nematocysts that are
connected with the umbrella margin by a line. Ocelli
and statocysts absent. Tentacles absent in P. nbyssicoln.

Type Species: Proboscidactyla flnvicirrntn Brandt, 1835.

Remarks: The life cycles within this genus are now
rather well known (e.g.. Hand 1954; Brinckmann-Voss
& Vannucci 1965). Only one species is known from
New Zealand, based on juveniles.

Proboscidactyla sp. (Fig. 52a-b)

Material Examined:

About 30 medusae collected by T. Barnett, June to August
1983 and February to July 1984, Leigh Marine Reserve,
some deposited.

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Fig. 52. Proboscidactyla sp., from preserved samples, a)
medusa with medusae buds; scale bar 02 min. b) nemato-
cysts: macrobasic eurytele from exumbrella, from tentacles,
discharged eurytele, anisorhiza, discharged anisorhiza; scale
bar 10 pm.

Description: Medusa up to 1 mm high, wider than
high, bell almost hemispherical, jelly moderately thick,
slightly thicker at apex. Velum spans approximately
one-fifth of radius. Exumbrella bears 4 interradial
clusters of nematocysts from where a fine line leads
to the umbrella margin. About 12 nematocyst capsules
per cluster. Manubrium about half of the height of bell
cavity, mouth either simple or with 4 irregular per-
radial lips, mouth rim with nematocysts. Medusae

buds arise from manubrium base at origin of radial
canals, with short stem but not on blastostyles. Four
radial canals and a very thin, solid circular strand
present. With 4 large perradial marginal bulbs with
black pigment granules. Four perradial tentacles,
longer than bell height, tapering, with evenly dis¬
tributed nematocysts. No ocelli present. No gonads
present. Nematocysts:

a) macrobasic euryteles from exumbrella, (15-
18) x (10-11) pm, s = 6.9.

b) macrobasic euryteles in tentacles, (9-11) x (6-7) pm,
s - 7.6.

c) merotrichous anisorhizas, in tentacles, spines very
fine, (4-5) x (3.5-4) pm.

Remarks: This Proboscidactyla species was found in very
large numbers by T. Barnett (1985, as unknown Tubu-
lariidae sp. A.) during autumn and winter months.
The medusa resembles some juvenile forms of Probo¬
scidactyla ornata, especially the figure of P. ornata var.
stolonifera given in Hartlaub (1918, fig. 322). Probo¬
scidactyla ornata as understood at present is an
extremely variable species and it may well be that
several species have been lumped together (cf. Mayer
1910; Hartlaub 1918; Kramp 1961). The medusae from
Leigh differed from young P. ornata in lacking
desmonemes and in developing their medusae buds
not on blastostyles but directly on the base of the manu¬
brium. Brinckmann-Voss and Vannucci (1965) were
able to reveal the life cycle of P. ornata and found that
juvenile medusae also had desmonemes. Although not
all nematocysts may have been found in the preserved
samples available, the medusae from New Zealand
probably are referrable to a new species. As only im¬
mature animals were found and the adult morphology
in Proboscidach/la is considerably different, it seems
advisable to defer creation of a new species name until
the life cycle is known.

Records from New Zealand: Leigh Marine Reserve
(Barnett 1985).

Family EUCODONIIDAE n. fam.

Anthomedusae without pointed apical projection.
Exumbrella without nematocyst tracks. Manubrium
on peduncle, quadrangular, with medusae budding
from stomach. Gonads completely surrounding
stomach. Four radial canals and circular canal, no
mesenteries. Four perradial marginal bulbs and four
tentacles; tentacles with terminal swelling. No ocelli.
Nematocysts are microbasic euryteles and desmo¬
nemes.

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Remarks: The family comprises only the genus Euco-
iionium. The systematic position of this monotypic
family and genus is discussed below.

Eucodonium Hartlaub, 1907

As in the family diagnosis.

Type Species: Eucodonium brownei Hartlaub, 1907.

Remarks: Eucodonium includes only E. broumei. The only
other described species, E. nrcticn Hand & Kan, 1961,
was synonymised with Plotocnide borealis Wagner, 1885
by Arai and Brinckmann-Voss (1980).

Eucodonium brownei Hartlaub, 1907 (Fig. 53a-b)

Dipurena sp.: Browne 1896: 473, pi. 16, fig. 2.

Eucodonium brownei Hartlaub 1907: 71, fig. 67; Russell 1953:
93, fig. 40; Vannucci 1957: 43, figs 2-3; Kramp 1959: 91,
fig. 44; Kramp 1961: 36; Brinckmann-Voss 1970: 19, pi.
2, fig. 4, text-figs 16-19; Petersen 1990: 217.

Material Examined:

14 medusae collected by T. Barnett from Leigh Marine
Reserve and Whangateau Harbour, 1980 and 1984,
February to March, all infertile, three cleared with lactic
acid for closer anatomical and nematocyst examinations,
some material deposited.

Descript ion: Medusa bell-shaped, up to 1 mm high and
as broad or slightly less broad, lateral walls thin, apex
with thickened jelly. Manubrium on a broad, well-
developed peduncle. Manubrium cylindrical, half as
long as bell cavity, mouth quadrangular with 4
inconspicuous perradial lips, each containing a group
of about 100 nematocysts. Medusae buds arise from
middle region of stomach. Four very narrow radial
canals and circular canal present. Four small marginal
bulbs containing blackish pigment granules. With four
equally developed tentacles with a conspicuous
terminal swelling. Tentacle gastrodermis chordoid.
Terminal swellings with swollen gastrodermis and
epidermis cells with fibrous structure. Nematocysts
not only present in terminal swelling but also along
the tentacles. No ocelli present. Nematocysts:

a) microbasic euryteles from tentacles, (7-8.5) x(3.5-
4.5) pm, s = 0.9.

b) heteronemes (microbasic euryteles?) from lips, (5-
6) x (2-3) pm.

c) desmonemes from tentacles, (4-5) x (2-3.5) pm.
Type Locality: Plymouth, England.

Fig. 53. Eucodonium broumei from preserved samples from
Leigh, a) medusa with buds on manubrium; scale bar
0.2 mm. b) nematocysts: microbasic eurytele from tentacle,
heteroneme from manubrium, desmoneme; scale bar 10 pm.

Remarks: The medusae from Leigh agreed well with
other descriptions of Eucodonium brownei, although the
terminal swelling of the tentacles was not as large as
depicted by some authors. According to Brinckmann-
Voss (1970), the gonads encircle the manubrium
completely in mature animals and there is no further
medusa budding. The mouth has been described by
early authors as simple and tubular, but Brinckmann-
Voss (1970) corrected this, observing four inconspicu¬
ous lips, also seen in the present samples. Closer micro¬
scopic examination revealed that the tips of these lips
contain nematocysts. There are thus four perradial
groups of nematocysts, separated in the interradii. The
nematocysts are rather deeply embedded in the lips
and do not protrude as in Podocoryna medusae, thus
are not easily seen at low magnifications. Contrary to
Brinckmann-Voss (1970), no stenoteles could be found
in three thoroughly examined animals. Picard (1955)
also found only euryteles and desmonemes, and the

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report of stenoteles for E. broumei may be considered
erroneous. At present, E. broumei cannot be placed in
any known family. Previously this species has been
associated either with the Tubulariidae, Corymor-
phidae, or placed as Capitata incertae sedis (Kramp
1961; Brinckmann-Voss 1970; Petersen 1990). All these
groups must be ruled out as E. broumei does not have
stenoteles and thus should not be placed among the
Capitata (cf. Picard 1955). The four perradial nemato-
cyst clusters embedded in the mouth margin ally E.
broumei closer to genera like Podocoryna and Cytaeis,
but still without corresponding to them. For these
reasons the new family Eucodoniidae is established
here to accommodate E. broumei.

Barnett (1985) found E. broumei in comparatively
high numbers during summer months. The species has
not so far been found in the wider Pacific, and may
have been introduced recently through human
activities. Brinckmann-Voss (1970) also indicated that
E. broumei occurs only during summer months in the
northern hemisphere. In the laboratory, lowering of
temperature causes a cessation of medusae budding,
regardless of season, which can be seen as a pre¬
condition for gonad maturation.

Records from New Zealand: Leigh Marine Reserve,
Whangateau Harbour (Barnett 1985),

Other Records: Northeastern Atlantic; Mediterranean;
Brazil (Kramp 1961).

Order CAPITATA

Athecate hydroids typically having capitate tentacles.
In some species these present only during develop¬
ment, in some only moniliform or even only filiform
tentacles present. Medusae typical single antho-
medusae, not colonial. Cnidome always includes
stenoteles in at least one stage.

Remarks: According to Petersen (1990), the syn-
apomorphy uniting the Capitata is moniliform
tentacles. These have, however, been modified very
early to either capitate tentacles or reverted to filiform
tentacles again. Despite this difficulty, at present it is
rather easy to recognise members of the Capitata.
Hydroids or solitary Anthomedusae possessing
stenoteles are always included in the Capitata. The
presence of stenoteles is a good diagnostic tool, but
most probably a plesiomorphic character because
other groups (Trachymedusae, Cubozoa) also have
them.

Family POLYORCHIDAE Agassiz, 1862

Medusae with gastric peduncle. Stomach prismatic
the four oral lips crowded with nematocysts. With
four sac-shaped or spiral, or several sausage-shaped
stomach pouches on peduncle only. Gonads sur¬
rounding stomach pouches. Four radial canals, with
or without blind side branches. Four to 260 tentacles
with stout, elongate bulbs with abaxial ocelli. Hydroids
unknown (after Petersen 1990).

Remarks: The polyps of this family are still unknown.
The putative hydroid of Polyorchis penicillatus was
described by Brinckmann-Voss (1977), but her work
was based on an uncertain linkage of life stages
(Brinckmann-Voss, pers. comm.).

Only Tiaricodon is known from New Zealand.

Tiaricodon Browne, 1902

Medusa with four perradial, imperfectly moniliform
tentacles with stout, elongated bulbs surrounded by
thickened, nematocyst-studded epidermis. With
abaxial ocelli. Stomach with short, sac-like perradial
lobes; mouth with four distinct, frilled lips with band
of nematocysts. Gonads on surface of stomach and on
stomach lobes. Four radial canals without diverticulae.

Type Species: Tiaricodon coeruleus Browne, 1902.

Remarks: Tiaricodon was previously allied with the
family Moerisiidae. Petersen (1990) referred it to the
family Polyorchidae and his arguments are followed
here. Only T. coeruleus is known in this genus. The
polyp stage is unknown.

Tiaricodon sp. (Fig. 54a-d

Material Examined:

1 young medusa collected 15.6.1994 in plankton Evans Bay
height 2.6 mm, infertile, cultivated until 19.8.1994, fed daily
with several Artemia nauplii, maximal height attained
7 mm, preserved after microbial infection caused deteri¬
oration, deposited.

4 T. coeruleus from Stanley Harbour, Falkland Islands, Soutli
Atlantic, collected by Vallentin 17.3.1902, held by ZMC.
identified by P. Kramp, size up to 18 mm, some mature
(figured in 54e).

1 T. coendeus from Valparaiso Bay, South America, collected
15.9.1958 by E. Fagetti, identified by P. Kramp, 5 mm high
but shrunken, stomach sacs present, immature.

Description

Young medusa: Youngest observed stage 2.6 mm high.

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i >4. a) to d) Tiaricodon sp., all from life, a) medusa found in the plankton of Evans Bay; scale bar 1 mm. b) same medusa
xanr 1 month, same scale as a), c) Nematocysts, from top: stenotele, same discharged, desmoneme, same discharged; scale
' 10 jim. d) lateral view of tentacle bulbs showing nematocyst ring (arrow). Note the ocellus on the abaxial spur of the
^rxade base; cc circular canal, ex exumbrella, rc radial canal, te tentacles; scale bar 0.2 mm. e) Tiaricodon coendeus from the
-ykland Islands, South Atlantic; scale bar 2 mm.

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bell-shaped with parallel lateral walls and conical top,
with very slight gastric peduncle. Exumbrella with
scattered stenoteles, denser towards margin. With ring
of thickened epidermis containing nematocysts
beneath circular canal. Manubrium tubular, reaching
almost the level of the velum, cylindrical with quad¬
rangular base, lacking stomach pouches. Four radial
canals and a circular canal, all thin and smooth. Radial
canals without dilatation on entering circular canal.
Four tentacles, contracted ~1.5 times bell length, length
when expanded more than three times bell size.
Tentacles with base free of nematocyst clusters and
an abaxial process adhering on exumbrella. This spur
bears a large ocellus. Tentacles hollow, with many
nematocyst clasps in indistinct spirals and a larger
terminal cluster. Radial and ring canal with golden
colour through zooxanthellae. Nematocysts:

a) stenoteles, (18-20.5) x (13.5-15.5) pm.

b) desmonemes, discharged with four coils, (10.5-
15.5) x (4.5-6.5) pm.

c) heteronemes.

Premature medusa: Medusa 7 mm high and 4 mm
wide. Umbrella bell-shaped with rounded apex, jelly
thick, at apex three times as thick as lateral walls. With
shallow gastric peduncle. Epidermal ring beneath
circular canal not observable. Manubrium almost as
long as bell cavity, with prismatic stomach, mouth with
4 perradial lips. Mouth margin thickened with
nematocysts. Base of stomach cruciform, with small
pouches that also differ in colour from radial canals.
Incipient gonad tissue (oocytes) visible on stomach
pouches. Tentacles thick, tapering, with bullet-shaped
terminal cluster. Ocelli on abaxial process of tentacle
bulb.

Remarks: Only one young medusa of Tiaricodon sp.
was found in Wellington Harbour during winter. After
several weeks of cultivation (Fig. 54b) the basal sacs
started to grow and some small oocytes began to
develop. Comparison with Tiaricodon coerideas from
the type locality showed that the medusa from Well¬
ington most probably belongs to this species (cf, Fig.
54e), but because the observed animal from Wellington
was immature, it was not definitely assigned to T. coe-
rideas. Only if mature animals are discovered in New
Zealand can the present observation be used as a
record for this species.

The basal sacs and especially the abaxial process
of the tentacle base bearing the ocellus (Figs 54b, d, e)
are very characteristic of Tiaricodon coeruleus . Contrary
to earlier descriptions, T. coeruleus from the type
locality does not always have a peduncle or has only
a very indistinct one.

In the youngest stage obtained from Wellington

(Fig. 54a, d) there was a ring of thickened epidermis
containing nematocysts beneath the radial canal. This
ring is most probably reduced during later develop¬
ment. Adult T. coeruleus from South America do not
have it.

Tiaricodon sp. is a very active swimmer, consumin
large numbers of Artemia nauplii (5-10 per day
Despite this, it grew only very slowly to a size of 7 mm.
after which it was preserved because it started to
deteriorate.

Records from New Zealand: Wellington Harbour (this
study).

Family ZANCLEIDAE Russell, 1953

Colonial hydroids with claviform or cylindrical
hydranths. Hydranths monomorphic or polymorphic.
With one oral whorl of four to six capitate tentacles
and below them numerous scattered capitate or
moniliform tentacles; rarely tentacles reduced.
Hydrocaulus short, unbranched, not clearly demar¬
cated from hydranth, arising from creeping stolons.
Perisarc covering stolons and it may form a low cup
around base of hydrocaulus, or covering hydrocaulus
entirely. Gonophores arise on short pedicels or in
clusters on short-branched blastostyles either among
tentacles, below tentacles, or from stolons. Gono¬
phores are liberated as free medusae with evenly
rounded umbrella. With exumbrellar nematocysts
confined to perradial patches of specialised tissue.
With either two or four solid marginal tentacles with
abaxial stalked cnidophores or tentacles lacking. Ocelli
absent. Gonads usually differentiated in interradial
position, rarely in a single mass around manubrium
of eumedusoids. Cnidome with stenoteles and
normally with macrobasic heteronemes, desmonemes
absent.

Remarks: The above diagnosis was taken from Petersen
(1990) and modified according to Boero and Hewitt
(1992) to accommodate polymorphic polyps and Halo-
coryne epizoica Hadzi, 1917. According to Petersen
(1990), the family comprises Asyncoryne and Z anclea
sensu lato. The monotypic genera Ctenaria and Oonautes
are not included. Ctenaria ctenophora Haeckel, 1879 is
here considered a doubtful species, and Oonautes
hanseni Damas, 1936 is of uncertain affinity. Both
species are normally included in the Zancleidae (e.g.,
Kramp 1961; Bouilon 1985b), but they are better
excluded as incertae sedis. Only Zanclea is known from
New Zealand.

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Zanclea Gegenbaur, 1857 emend.

Haloconjne Hadzi, 1917 new synonym.

Zanclella Roero & Hewitt, 1992 new synonym,
for more see Calder (1988)

Colonial hydroids arising from ramified stolons. Often
associated with bryozoans. Hydranths monomorphic
or polymorphic, with elongated cylindrical or fusi¬
form body, with an oral whorl of 1 to 6 capitate ten¬
tacles and normally additional capitate tentacles
scattered over the hydranth body. Tentacles rarely
reduced to mere nematocyst patches. Stolons covered
by perisarc which can extend up to the hydrocaulus.
Medusae-buds arise either scattered among or under
tentacles or from stolons. Gonophores develop singly
on short pedicels or on short, branching blastostyles.
Medusae with two or four perradial tentacles with
abaxial stalked cnidophores or tentacles absent. Ocelli
lacking. With nematocyst clusters on exumbrella
above tentacle bulbs that contain stenoteles. Nemato-
cysts of medusa includes stenoteles and macrobasic
heteronemes.

Type Species: Zanclea costata Gegenbaur, 1857.

Remarks: The diagnosis of the genus Zanclea is
emended here to include Z. polyniorphn n.sp. This
species has polymorphic hydranths and an obligatory
association with bryozoans as in Zmiclella Boero &
Hewitt, 1992. It does not conform to Zmiclella , how¬
ever, because its polyps have normally developed
tentacles and the medusa agrees with the previously
accepted diagnosis of Zanclea. Polymorphic polyps and
association with bryozoans are apparent synapo-
morphies of a clade that includes the present new
species and Zanclella bnjozoophila. It is therefore not
possible to widen the genus definition of Zanclea to
include only the new species as this would render
Zanclea obviously paraphyletic. Because no apparent
synapomorphies are available for the remaining
species with monomorphic polyps, they are thus
probably paraphyletic. The generic diagnosis of Zanclea
is therefore widened to accommodate not only the new
species but also Zanclella .

The monotypic genus Haloconjne Hadzi, 1917 was
shown by Piraino et al (1992) and Boero and Hewitt
(1992) to be closely related to Zanclella. It shares with
the latter polymorphic polyps, an obligatory associ¬
ation with bryozoans, a reduction of tentacles in the
medusa, and gonads encircling the manubrium. All
these characters are synapomorphies within the genus
as defined here. Haloconjne is therefore also
incorporated into the new diagnosis of Zanclea to avoid
paraphyletic taxa. Zanclella and Halocoryne thus

become synonyms of Zanclea.

Only one species of Zanclea is known in New Zea¬
land.

Zanclea polymoiyha n.sp. (Figs 55a-g, 56a-b)

Material Examine*:

1 colony collected east of NIWA buildings, Greta Point,
Wellington, under rocks, intertidal, 17.11.1993, infertile.
Ca. 12 colonies from type locality, collected 17.10.1994 and
2.11.1994, 0.5 m, infertile at date of collection, all on the
bryozoan Rhynchozoon larreyi (kindly identified by Dr D.
Gordon, see also Gordon & Mawatari 1992), several
colonies cultivated in running seawater and fed every
second day with Arte min nauplii. Many new stolons grew
between zooids but never extended beyond the bryozoan
host. After 2-4 weeks medusae buds appeared and about
30 free medusae from 3 colonies were obtained. Medusae
were cultivated at room temperature and fed daily with
Artemia parts. After 7 days gonads (only males seen)
became mature (as judged by their turning opaque). Many
medusae were lost during cultivation as they are prone to
microbial infections. After 8 days the medusae looked very
closely like medusae from the plankton, only being smaller
in size (1 mm) and the apical thickening was not as thick
as seen in medusae from the plankton. One hydroid colony
deposited as holotype H-653, young medusae deposited
as paratype P-1084.

8 medusae, collected 14.4.1994 bn the surface plankton near
Seatoun, several mature which spawned later on, some
deposited as paratype P-1084.

7 preserved medusae collected by T. Barnett, Leigh Marine
Reserve, 1983-1984, labelled Zanclea costata.

Description

Polyp stage: Hydroid colonies growing within and on
colonies of Rhynchozoon larreyi (Audouin, 1826)
(Bryozoa, Cheilostomatida). Polyps polymorphic,
sessile, without marked caulus, arising from stolons
which are mostly embedded in the bryozoan colony
or covered by calcified material secreted by the
bryozoans. The hydranths cannot completely with¬
draw into the bryozoan colony and are always visible.
Stolons may be visible in wild colonies only near the
edge of the bryozoan colony where they run along the
margins of the zooids. Stolons are covered by a thin,
single-layered perisarc, there is no perisarc cup present
at the base of the polyps.

Polyps differentiate into gastrozooids, dactylo-
zooids, and rarely into gonozooids. Gastrozooids are
normally 1 mm high but can reach 2 mm, claviform
with a thicker distal end and a white oral region and
orange body. With 4-6 capitate oral tentacles and 8-
14 scattered capitate tentacles below them. Tentacles
become progressively shorter towards hydranth base.
Dactylozooids are very long and thin, extensible for

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Fig. 55. Zanclea polymoiplta n.sp., all from life, a) fertile colony on its bryozoan host Rhynchozoon Intrey i; scale bar 0.5 mm.
b) gastrozooid, same scale as d). c) dactylozooid, same scale as d). d) Medusae buds arising from stolons (left) or gonozooid
like polyps (right); scale bar 0.2 mm. e) nematocysts of polyp stage, anticlockwise: macrobasic eurytele, same discharged,
shaft of the latter, small stenotele, larger stenotele; scale bar 10 Jim. f) newly released medusa; scale bar 0.1 mm. g) morphology
of tentacle with cnidophore; scale bar 20 Jim.

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56. Zanclea polymoij)ha n.sp. from life, a) adult medusa
plankton; scale bar 0.25 mm. b) nematocysts of medusa
larger stenotele, smaller stenotele, macrobasic eury-
same discharged; scale bar 10 pm.

' to 5 mm, more frequently present near margin of
zoan colony, with oral region swollen and hypo-
Tie white, with 1-4 capitate oral tentacles and 0-4
capitate tentacles widely scattered on body. At
ast some of the dactylozooids still have a mouth and
able to consume small prey, but mostly prey caught
' dactylozooids is eaten by neighbouring gastro-
- > nds.

Gonophores normally arise from stolons, either
mgly on stems or from short blastostyles that branch
or twice. Occasionally gonophores are produced

below the tentacle zone of gastrozooid-like polyps that
have a reduced number of tentacles. Gonophores
develop into free medusae. The buds soon become
bilaterally symmetrical by the pronounced growth of
two opposite marginal bulbs. Nematocysts:

a) macrobasic euryteles, concentrated near mouth,
with three bands of spines spiralling around distal
quarter of shaft, (22-25) x (10.5-12) pm, s = 8-10.

b) smaller stenoteles, (7.5-8.5) x (5-6) pm,

c) larger stenoteles, (12.5-14.5) x (10-11) pm.

Young medusa: Newly released medusa bell-shaped,
not bilaterally flattened, as broad as high or slightly
higher, approximately 0.6 mm high, jelly evenly thin.
With 4 perradial exumbrellar ridges extending from
bell margin upwards one-third of bell, each bearing
an oval to fusiform patch of nematocytes with steno¬
teles. Occasionally some single nematocysts on
exumbrella. Velum spanning approximately half of
radius. Manubrium length half of bell height, simple,
tubular with circular mouth. Four incipient interradial
gonads visible. With 4 radial canals without median
swelling, ending in 2 opposite marginal bulbs with
tentacles and 2 small bulbs without tentacles. Circular
canal thinner than radial canals. One pair of opposite
tentacles, extended somewhat longer than bell height,
rapidly tapering from proximal to distal, with around
70 abaxial cnidophores on long contractile stalks.
Cnidophores and stalks are purely epidermal and
contain 3-4 euryteles. Tentacle main axis additionally
with many protruding cell processes. No ocelli present.
Nematocysts:

a) macrobasic euryteles, undischarged capsule often
with a concave side, shaft not coiled, few spines
on everted shaft, (7.5-9) x (5-6) pm, s = 3.5-5.4.

b) smaller stenoteles, (6-9) x (4-6.5) pm.

c) larger stenoteles, (13-15) x (10-12) pm.

Mature medusa: Similar to young medusa, only larger
(1-1.6 mm), jelly thicker. Apex mostly with a thickened
jelly of variable height, up to approximately 2.5 times
the thickness of lateral walls. The exumbrellar
nematocyst patches are reduced to narrow bands. The
tentacles are longer and with many more cnidophores.
The gonads are 4 bulging pads in interradial positions
on the manubrium. Nematocysts:

a) macrobasic euryteles, (8.5-11) x (5-6.5) pm, s = 3.3-
4.3.

b) smaller stenoteles, (8-10.5) x (6-8.5) pm.

c) larger stenoteles, (12-16) x (9.5-13) pm.

Egg size: 99 mm (s.d. 5 mm, n = 8).

Type Locality South of NIWA buildings, Greta Point,
Wellington Harbour, depth 0.5 m, on colonies of
Rhynchozoon larreyi (Bryozoa) on the underside of loose
bricks.

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Etymology: The species name refers to the polymorphic
polyps.

Remarks: With its polymorphic polyps Znnclen
polymorphn somewhat resembles Znnclelln Boero &
Hewitt, 1992, although the latter has more-reduced
polyps. The medusa of Z. polymoiyhn is also different
in having two fully developed tentacles. The medusa
alone is indistinguishable from medusae commonly
referred to Znnclen costntn (cf. Russell & Rees 1936;
Russell 1953; Kramp 1968; Brinckmann-Voss 1970).
The systematics of this species is very complicated and
unsettled. Some authors have already indicated that
more than one species may be lumped together under
the name Z. costntn (cf. Berhaut 1969; Millard & Bouil¬
lon 1973; Tregouboff & Rose 1978). With Z. polymorphn
the uncertainty becomes even larger. It seems that the
various Z. costntn- like medusae are not properly
identifiable if only this stage is known, and complete
life-cycle information will be necessary for all future
identification. As shown by Bouillon (1978b) there are
also several valid Znnclen species known from their
medusa stage only. This indicates that there are a large
number of Znnclen species to be properly investigated
and described.

Znnclen polymorphn polyps fed well on Artemin
nauplii, but all colonies degenerated after few weeks,
perhaps because the bryozoan was not surviving.
Albeit with difficulty, several released medusae could
be cultivated until their gonads appeared mature. They
reached a size of about 1 mm and differed from the
medusae found in the plankton only in having a less
thickened apical jelly. This can most probably be
attributed to the cultivation conditions and younger
age, however, as similar effects were observed with
other medusae (pers. obs.). The apical thickening is
also rather variable in medusae from the plankton.
There were no differences in variety or size of nemato-
cysts between the cultivated and wild medusae,
therefore the Znnclen medusae from the plankton of
Wellington Harbour can quite reliably be attributed
to Z. polymorphn . No other zancleid was found and Z.
polymorphn polyps were quite abundant at the type
locality. The medusae originating from Leigh are not
distinguishable from the ones found in Wellington and
were tentatively assigned to the same species.

The bryozoan host Rhynchozoon Inrreyi is abundant
throughout New Zealand (Gordon & Mawatari 1992).
Not all colonies host Z. polymorphn, however, and the
hydroid was never found on other bryozoans despite
a large number of examined species. At the type
locality only about one-third of the R. Inrreyi colonies
encrusting the underside of stones had visible polyps.
Colonies of R. Inrreyi on gastropod or bivalve shells,
or on seaweeds did not host the hydroid. Gordon and

Mawatari (1992, fig. 3b) depicted a R. Inrreyi colony
originating from South Island (D.P. Gordon pers
comm.) which clearly shows a similar hydroid like Z.
polymorphn and most presumably also belongs to this
species.

Rhynchozoon Inrreyi is also widespread outside New
Zealand (Turkey, Red Sea, Indonesia, Lord Howe
Island, Victoria, see Gordon & Mawatari 1992). In
colonies from the Red Sea it is known to host a hydroid
of the genus Znnclen in almost all examined colonie-
and this association may contribute to the com¬
petitiveness of the bryozoan (Ristedt & Schuhmacher
1985). Whether it hosts the same species of Znnclen as
in New Zealand cannot be decided from the infor¬
mation available. Rhynchozoon Inrreyi has a frontal
budding pattern resulting in thick colonies with a
complicated and highly structured surface. The
microrelief may offer the hydroids protection from
abrasion and grazing as they can withdraw into the
cavities and depressions.

Records from New Zealand: Wellington Harbour
?South Island (Gordon and Mawatari 1992), Leigh
Marine Reserve (Barnett 1985, as Znnclen costntn).

Family CLADOCORYNIDAE Allman, 1872

Hydroids with club-shaped hydranths, moniliform or
capitate tentacles in one oral whorl and moniliform
or modified moniliform aboral tentacles scattered or
in whorls. With rounded nematocyst patches on body
wall below oral tentacles. Stems simple or sparingly
branched, arising from attached stolons. Gonophores
develop singly or on short, branched blastostyles on
lower part of hydranth. Gonophores either liberated
as free medusae or remaining fixed as sporosacs.
Newly liberated medusa with two perradial tentacles
with stalked cnidophores and two non-tentacular
bulbs; exumbrellar nematocyst pouches absent.
Cnidome of both hydroid and medusa comprising
macrobasic euryteles and stenoteles (Petersen 1990).

Remarks: This family comprises the genera Pteroclnva
and Clndoconjne. Only the latter genus is known from
New Zealand.

Cladoconjne Rotch, 1871
Synonyms: see Petersen (1990).

Hydroid colonies with club-shaped hydranths with
an oral whorl of four to six short, capitate tentacles
and one to four whorls of branched-capitate aboral

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tentacles. One or two series of oval patches of
macrobasic euryteles on hydranth body. Hydrocaulus
ong, unbranched or sparingly branched, covered by
r*erisarc. Cauli arising from attached stolons. Gono-
shores carried singly on short pedicels between or over
aboral tentacles. Gonophores remain fixed as sporo-
sacs in the presently known species (after Petersen
1990).

Type Species: Cladocoryne floccosa Rotch, 1871.

Remarks: Only the type species is known from New

Zealand.

Cladoconjne floccosa Rotch, 1871 (Fig. 57a-c)

'+kk:ojynefloccosn Rotch, 1871: 228; Brinckmann-Voss 1970:
69, figs 80-82; Millard 1975: 65, figs 21A-B; Bouillon et
(d. 1987: 297, figs 1 & 5-6; Hirohito 1988: 52, figs 16b-f.

Iaterial Examined:

live, infertile colony with 6 hydranths, on holdfasts of the
alga Macrocystis pyrifera, 10.3.1994, drift, Greta Point,
Wellington.

Descriphon: Colonial hydroids reaching several milli¬
metres in height, arising from ramified stolons. Stems
not branched. Stems covered by perisarc, smooth or
with annulated stretches and becoming thinner distally
to terminate below hydranth. Hydranth cylindrical,
up to 1 mm high, with dome-shaped hypostome, with
a single whorl of 4 capitate oral tentacles and 3 alter¬
nating whorls of aboral tentacles. Aboral tentacles
branched, side-branches short, in 2 lateral rows and 1
median row on upper side. Lateral rows with up to 7
tentacles, median rows with 0-2 tentacles. All side
branches and end of main branch with a terminal,
spherical nematocyst cluster. Colour white.
Nematocysts:

a) telotrichous macrobasic euryteles, on body wall
between oral tentacles and most proximal aboral
tentacles, (33-41) x (16-17) pm, s > 10.

b) smaller stenoteles, (6-7) x (4-5) pm.

c) medium sized stenoteles, (9) x (6-7) pm.

d) larger stenoteles, (15-16.5) x (12-13.5) pm.

Type Locality: Herm, Channel Islands, United King¬
dom.

Fig. 57. Cladocorytte floccosa, from life, a) distal part ot cauius and nydrantti; scale bar 0.2 mm. b) terminal part of an aboral
wtacle showing branching, CL; scale bar 50 pm. c) nematocysts: column of stenoteles, macrobasic eurytele, same discharged;
■cale bar 10 pm.

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Remarks: According to Millard (1975), the gonophores
are borne on the hydranth among the aboral tentacles
on short pedicels; they are spherical, reaching 0.34 mm
in diameter, remaining fixed as sporosacs of the
cryptomedusoid type.

Although the examined colonies were not fertile,
the identification of C. floccosa is rather unambiguous
as it had three aboral tentacle whorls (cf. Bouillon et
ai 1987). The macrobasic euryteles are clustered in
groups on the body wall, but are not easy to see under
a dissecting microscope.

Records from New Zealand: Wellington (new record).

Distribution: Circumglobal in tropical and subtropical
waters, occasionally extending into temperate areas
(Millard 1975).

Family PORPITIDAE Goldfuss, 1818

Hydroids developed as floating colony. Underside
with a large central gastrozooid surrounded by
gonozooids and peripheral dactylozooids. Float con¬
tains a perisarc disc with concentric air chambers and
a complex system of gastrodermal canals. Gono¬
phores develop on basal parts of gonozooids and are
liberated as medusae. Umbrella of medusa with four
or eight tracks of nematocysts originating from mar¬
ginal bulbs. With four or eight radial canals and a ring
canal. Manubrium short, with circular mouth. Gonads
perradial or interradial; may be irregularly developed
so that number may vary. With two opposite marginal
tentacles terminating in large spherical nematocyst
knob. With or without an additional adaxially oriented
short tentacle developed from bulbs with tentacles.
Two or six marginal bulbs without tentacles. No ocelli
present. Zooxanthellae often present.

Remarks: Only Porpita and Velella are known from New
Zealand. Diagnostic characters:

Poiyita : float without sail, smooth surface, dactylo¬
zooids with capitate tentacles.

Velella : float with upright sail, dactylozooids without
tentacles.

Poipita Lamarck, 1801

Porpitid hydroids with disc-shaped float and mantle.
Float flat or with central bulge, without sail. Dactylo¬
zooids with three vertical rows of short, capitate
tentacles. Medusa with small manubrium. Juvenile
specimens without marginal tentacles; adults with one
to two slender, decidedly capitate tentacles. With eight

radial canals, their gastrodermal cells bearing algal
symbionts.

Type Species: Poipita poipita (Linnaeus, 1758).

Remarks: Only the type species is known from Nev>
Zealand.

Poipita poipita (Linnaeus, 1758) (Fig. 58a-b

Medusa poipita Linnaeus, 1758: 659.

Porpita paciifica Lesson, 1826: pi. 7, figs 3, 3'.

Porpita umbella : Vanhoeffen 1906: 39, figs 64-65.

Porpita pacifica : Powell 1947: 5, figure.

Porpita poipita: Brinckman n Voss 1970: 38, figs 40-42; Danid
1976: 111, fig. la-d; Calder 1988: 77, figs 65-67 (cum sy«.
Pages et al 1992: 23, fig. 22.

Fig. 58. Porpita porpita. a) polyp stage, drawn from
photograph, no scale available, b) mature medusa from
plankton, redrawn from Bouillon (1984) with permission
of the publisher. Note: the medusa is not yet known from
New Zealand; scale bar 1 mm.

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Material Examined:

\ T o material from New Zealand seen.

Description:

Polyp stage (after Vanhoeffen 1906; Calder 1988):
Hydroid colony floating on water surface, diameter
up to 30 mm, mostly smaller, with disc-shaped mantle
and internal float; upper surface slightly convex, with
a central pore and numerous stigmata. Mantle with
radiating gastrodermal canals; margin soft, flexible;
central region firm, with an internal chitinous float
consisting of a series of concentric chambers; a disc¬
shaped reservoir of nematocysts lying between float
and central gastrozooid. Undersurface with one large
central gastrozooid, a median circle of gonozooids, and
a peripheral circle of dactylozooids. Central gastro¬
zooid short and broad with a terminal mouth, without
tentacles or prominent nematocyst clusters. Gono¬
zooids clavate, lacking tentacles but with prominent
nematocyst clusters scattered over body, medusae
develop near base in clusters. Dactylozooids with ten¬
tacles, with a distal whorl of 4 capitate tentacles, body
with varying number of short, small capitate tentacles
in 3 vertical rows.

■ ematocysts: atrichous isorhizas, haplonemes, three
types of stenoteles.

Young medusa (after Brinckmann-Voss 1970 and
Bouillon 1984): Bell-shaped, slightly higher than wide,
height 0.3 mm. With 4 exumbrellar rows of nemato¬
cysts. Manubrium very short. Four broad radial canals,
circular canal missing. Tentacles lacking. Gastro¬
dermal system without lumen, composed of large,
vacuolated cells. Gastrodermis of radial canals with
zooxanthellae.

Adult medusa (after Bouillon 1984, not known from
New Zealand): Up to 2.5 mm high and 2 mm diameter,
bell rather conical, with jelly of even thickness.
Exumbrella with 8 perradial tracks of nematocysts,
only one capsule wide. Manubrium length one-third
of bell cavity, conical, with circular, hardly visible
mouth. Gonads normally in four perradial masses on
manubrium, but 3-8 gonads may be present. With 8
large radial canals with zooxanthellae. Circular canal
present. With 8 little developed marginal bulbs. Only
2 opposite tentacles present. Tentacles long and ending
in a voluminous terminal swelling. Tentacles develop
unequally and are often of different length. Often only
one tentacle present. Ocelli absent. Nematocysts:

a) stenoteles, (24) x (23) |im.

b) telotrichous macrobasic euryteles, (23) x (17) jim,
s~7.

Type Locality: India.

Remarks: Porpita porpita is often found stranded
together with Velella velella, though Porpita is normally
only present in few numbers.

Records from New Zealand (polyp): Muriwai Beach
(Powell 1947); Tawharanui Peninsula (D. P. Gordon,
pers. comm.).

Distribution: Circumglobal in warmer waters (polyp
stage, Daniel 1976). Adult medusa known from Papua
New Guinea (Bouillon 1984).

Velella Lamarck, 1801

Floating hydroid colonies with an upright sail; with a
central gastrozooid, numerous feeding gonozooids,
and dactylozooids. Free medusa with exumbrellar
nematocyst rows, with two pairs of opposite, perradial
tentacles, each tentacle with a large terminal nemato¬
cyst cluster, two perradial marginal bulbs without
tentacles. Stomach with tubular mouth. Gonads in
male divided. Female with one egg. (Brinckmann-Voss
1970).

Type Species: Velella velella (Linnaeus, 1758).

Velella velella (Linnaeus, 1758) (Fig. 59a-f)

Medusa velelln Linnaeus, 1758: 660.

Velella spirans: Vanhoeffen 1906: 37, figs 58-60.

Velella velella: Brinckmann-Voss 1964: 327, figs 1-3; Edward
1966a: 283; Brinckmann-Voss 1970: 34, figs 36-39; Daniel
1976:118, fig. lm-n; Larson 1980: 183, fig. 1; Calder 1988:
81, figs 58-59 (cum si///.); Pages et al 1992: 21, fig. 21.

Material Examined:

2 colonies held by Portobello Marine Laboratory, collected
17.1.1961 near Tokomairo mouth (Otago); 3 and 5 cm long,
both right sailing (sail running from northeast to south¬
west, see Edwards 1966).

8 colonies held by Portobello Marine Laboratory, collected
27.3.1971 near East Cape, 1-3.5 cm, all right sailing,

37 colonies held by MoNZ, collected 5.5.1975 in Island Bay,
Wellington, 1.5-4 cm, all right sailing.

8 colonies from NZOI Stn U799,42°33.8 r S, 170*34,E, 2.8.1990,
2-20 mm, all right sailing.

9 colonies from NZOI Stn U796,42°34'S, 170°34.4'E, 6.8.1990,
3 -20 mm, all right sailing .

3 floats collected in December 1993 by C. Battershill on Piha
Beach, 1.8-2.3 cm, all right sailing.

Several live colonies collected 19.3.1994 on east coast of
Kapiti Island, all examined right sailing, medusae released
and cultivated.

2 preserved colonies collected by M. McLean 16.4.1994 near
Poor Knights Islands, 4-5 cm, one right and one left sailing
form.

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Description:

Polyp stage: Hydroid colonies floating on water
surface, with flattened oval float and a triangular sail.
There are two mirror images of the animal (left and
right sailing). Float and sail are kept rigid by a chitinous
support covered by mantle tissue. Margin of float soft
and flexible. Chitinous float oval to slightly S-shaped
with concentric air chambers. Mantle tissue with
network of gastrodermal canals. In centre of underside
a single large gastrozooid encircled by a band of
medusae producing gonozooids and a peripheral band
of dactylozooids. Central feeding zooid broadly oval
with an elongated hypostome, without tentacles or

medusae buds. Gonozooids spindle-shaped with a
swollen mouth region, lacking tentacles but with warts
of nematocyst clusters concentrated in distal half. On
proximal half of hydranth numerous medusae buds
growing in groups from short blastostyles. Dactylo¬
zooids long and tapering, oval in cross section with
nematocysts concentrated on the narrow sides, mouth
lacking. Nematocysts:

a) larger stenoteles, (17.5-19.5) x (15.5-17) pm.

b) medium sized stenoteles, (14-16) x (11-12) pm.

c) smaller stenoteles, (10.5-12.5) x (8-9.5) pm.

d) isorhizas with very fine spiral pattern on thread
almost like atrich, (7.5-8.5) x (3-4) pm.

Fig. 59. Velella velella, from preserved material except e) and f). a) view of a colony (right sailing); scale bar 3 mm. b)
dactylozooid; scale bar 1 mm. c) gonozooid with medusae buds, same scale as b). d) central gastrozooid; scale bar 2 mm. e)
nematocysts in pairs of native and discharged capsule: stenotele, isorhiza, macrobasic eurytele of medusa; scale bar 10 pm
f) newly released medusa, zooxanthellae shaded; scale bar 0.2 mm.

100

@®®C

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Colours: float deeply blue when alive, medusae buds
vellow-olive from symbiotic algae.

Young medusa (from live samples): Newly released
medusa bell-shaped, up to 1 mm high and 0.8 mm in
diameter. Manubrium short, conical, red colour often
broad radial canals ending in 4 broad marginal bulbs
connected by radial canal. No tentacles present in
examined material. Many zooxanthellae are found in
groups along the radial canals. During development
present. With 4 perradial rows of stenoteles on
exumbrella originating from the marginal bulbs. Four
the zooxanthellae were reduced and an opposite
tentacle pair grew, many macrobasic euryteles were
produced. Nematocysts:

a) stenoteles, ca. 20 x 15 pm.

b) telotrichous macrobasic euryteles, (14-19.5) x (10-

14) pm, s = 7.

Adult medusa (not known from NZ): Up to 2.8 mm
high and 2 mm wide, bell cylindrical, with flat top,
jelly evenly thin. Exumbrella with numerous papillae.
Manubrium conical, length half to two-thirds of bell
cavity. Gonads on manubrium as 4 longitudinal
swellings in perradial position. With 4 marginal bulbs.
Two opposite bulbs lack tentacles, and the other bulbs
have 2 tentacles each, a short stout adaxial capitate
tentacle, and a longer axially-directed capitate tentacle.
Tentacles with parenchymatic gastrodermis. On the
abaxial side of each tentaculate marginal bulb is a
triangular patch of about 50 stenoteles which extends
m an irregular line to the bell apex. On the abaxial
side of each atentaculate bulb 15-20 stenoteles forming
a vertical, irregular double row extending a short
distance from the bell margin, and continuing as an
uregular line to the bell apex. Marginal sense organs
absent (after Brinckmann-Voss 1964; Larson 1980).
Nematocysts: stenoteles, macrobasic euryteles.
Colour: umbrella dark brown due to zooxanthellae.

Remarks: The newly-released medusae observed in this
c tudy did not have rudiments of tentacles and also
lacked macrobasic euryteles. This does not agree with
descriptions of Brinckmann-Voss (1970). However, the
euryteles were formed shortly afterwards and in some
medusae rudiments of tentacles also became visible.
Unfortunately the medusae could not be cultivated
for longer than seven days.

Although Velella velella can occur in enormous
swarms spreading over hundreds of kilometres, and
each individual colony can release thousands of
medusae, the adult medusa has only rarely been
reported from nature (by Metschnikoff after Brinck¬
mann-Voss 1970; Larson 1980). Metschnikoff's medusa
was a female that had one large, red egg. Brinckmann-

Voss (1964) managed to cultivate male medusae to
maturity. More information on the development of the
colony can also be found in Brinckmann-Voss (1970).
Larson (1980) described the only fully mature animal
known from nature. The animal was intensely pig¬
mented by zooxanthellae which suggests that the
medusae normally occur near the water surface.

Almost all polyp colonies observed in this study
had their sail in 'Tight sailing" position (see Edwards
1966a). The prevalence of one form in one region may
be due to sorting by prevailing winds (cf. Edwards
1966a).

Records from New Zealand: North Island (Powell
1947), South Island (see under Material examined).

Distribution: Widely distributed in Atlantic, Medi¬
terranean, Indian Ocean, Pacific Ocean.

Family CORYMORPHIDAE Allman, 1872

Solitary hydroids with one whorl of moniliform or
capitate oral tentacles or several close-set whorls of
filiform oral tentacles. With one to three whorls of
moniliform or filiform aboral tentacles. Hydrocaulus
long, lower end pointed or rounded, hollow or more
or less filled with parenchymatic gastrodermis. With
short papillae or longer filaments composed of epi¬
dermis with a core of chordoid gastrodermis termi¬
nating in non-ciliated statocysts. Gonophores carried
singly or on blastostyles just above the aboral tentacle
whorl. Gonophores remain fixed as sporosacs or are
liberated as free medusae. Medusa with umbrella that
may or may not have an apical projection. Manubrium
mostly not extending beyond umbrella margin; mouth
simple, circular. With one to four hollow, usually
moniliform tentacles. Ocelli lacking. Development
from egg to polyp via encysted gastrula which
develops directly into young polyp without planula
stage. Cnidome includes both stenoteles and
desmonemes (after Petersen 1990).

Remarks: The family Corymorphidae as recently
redefined by Petersen (1990) comprises the genera
Euphysa, Siphonohydra, Gymnogonos , Corymorpha,
Branchiocerianthus and Fukciurahydra. It is here modi¬
fied slightly to allow the inclusion of medusae with
manubria longer than the bell cavity. This becomes
necessary with the inclusion of Euphysa problematica
(see p. 106). The more classical concept of this family
(e.g., Bouillon 1985a) also contained genera that are
either problematic (e.g., Plotocnide) or may be referred
to other taxa (e.g., Eucodonium , Ralpharia). In New Zea¬
land only Corymorpha and Euphysa are known.

101

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Characteristics:

Corymorpha : medusa with pointed apical projection,
one tentacle only.

Euphysn : medusa with evenly rounded umbrella,
tentacles all of the same structure.

Conjmorpha M. Sars, 1835

Synonyms: (after Petersen 1990)

Steenstrupia Forbes, 1846
Euphysora Maas, 1905
Vannuccia Brinckmann-Voss, 1967
Gotoea Uchida, 1927
Eugotoea Margulis, 1989

Solitary hydroids with more or less vasiform hydranth,
one or several closely set oral whorls of sixteen or more
moniliform or filiform tentacles and one aboral whorl
of sixteen or more long non-contractile filiform
tentacles. Gastrodermal diaphragm parenchymatic.
Hydrocaulus stout, covered by thin perisarc, filled
with parenchymatic gastrodermis with long peripheral
canals; aboral third of caulus with papillae and, more
aborally, rooting filaments. These have gastrodermal,
non-ciliated statocysts in the species investigated. With
or without asexual reproduction through constriction
of tissue from aboral end of hydrocaulus.

Gonophores develop on blastostyles arranged in a
whorl over aboral tentacles. Gonophores remain either
fixed as sporosacs or are released as free medusae.
Medusa apex dome-shaped or pointed. With three
short or rudimentary tentacles and one long tentacle
that differs not merely in size, but also in structure.
Manubrium thin-walled, sausage-shaped with flared
mouth rim, reaching to umbrella margin (after
Petersen 1990).

Type Species: Conjnioijfhn nutans M. Sars, 1835.

Remarks: The revised generic concept of Corymorpha
includes genera like Euphysora and Vannuccia (Petersen
1990). The genus is now mainly defined through its
polyp phase which offers better characters. Some of
the characters of the medusa given in Petersen's diag¬
nosis may be problematic. The flared mouth rim, for
example, could not be seen in the available preserved
material and could not be verified from other sources.
More observations concerning this point are needed.
Despite this, the system of Petersen (1990) is an
improvement leading towards a more natural classi-

Fig. 60. Corymorpha forbesii. a) medusa from Leigh; scale bar 0.2 mm. b) nematocysts of medusa: large stenotele, small
stenotele, desmoneme, heteroneme, haploneme, heterotrichous anisorhiza, same discharged; scale bar 10 pm. c) polyp stage,
redrawn from Brinckmann-Voss (1970) with permission of the publisher; size of polyp 10 mm. Note: the hydroid has not yet
been found in New Zealand.

102

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fication. Characteristics of species known from New
Zealand:

C, forbesii : medusa with one thick tentacle with a
terminal swelling, no apical process present
C. furcata : medusa with apical process, with three
smaller and one large tentacle with branching ends.
C. in ter media : medusa with pointed apical process, one
slender, moniliform tentacle.

Coiymoifha forbesii (Mayer, 1894) (Fig. 60a-c)

Hybocodon forbesii Mayer, 1894: 236, pi. 1, fig. 1; Mayer 1910:
42, pi. 1, fig. 8, pi. 2, fig. 3; Uchida 1927: 193, fig. 30;
Kramp 1959: 87, fig. 36; Kramp 1961: 42; Kramp 1968:
13, fig. 22.

Hybocodon forbessi : Nail* 1951: 50, pi. 1, fig. 1 [incorrect
spelling].

Vamiuccia forbesii : Brinckmann-Voss 1967:1, figs 1-6; Brinck-
mann-Voss 1970: 14, pi. 1, figs 1-2, text-figs 9-11;
Bouillon 1978a: 268, fig 8.3-4; Bouillon 1978b: 20.
Corymorpha forbesii : Petersen 1990: 149, fig. 16B.

Material Examined:

3 medusae from Leigh Marine Reserve, 1 medusa from
Whangateau Harbour (Leigh) obtained from T. Barnett.

Description:

Medusa stage: Adult medusa with bell-shaped
umbrella up to 1.8 mm high, higher than wide, with¬
out apical process. Jelly evenly thin. Bell margin
slightly oblique. Without exumbrellar tracks of
nematocysts. Manubrium cylindrical, length half to
two-thirds of bell height. Gonads encircle manubrium
for almost all its length. Four narrow radial canals and
circular canal present. One voluminous tentacle only.
Tentacle hollow for half its length, terminal region
except very distal end with a swollen epidermis con¬
taining many nematocysts. On perradial sites other
than occupied by tentacle only marginal bulbs present.
Bulb opposite tentacle slightly larger than the other
ones. Nematocysts:

a) larger stenoteles, rare, (18) x (15.5) pm.

b) smaller stenoteles, (6.5-8.5) x (4.5-5.5) pm.

c) desmonemes, (6-7.5) x (4.5-5.5) pm.

d) heterotrichous anisorhizas, thread tapering only
slowly, diameter (10-11.5) pm.

e) heteronemes (microbasic euryteles?, not seen dis¬
charged), (8) x (4) pm.

f) haplonemes (isorhizas?, not seen discharged), (7.5-
9) x (3.5-4) pm.

Polyp stage (after Brinckmann-Voss 1967, 1970, not
known from New Zealand): Solitary hydroids, up to
3 cm high, but length of caulus very variable. Polyp
composed of caulus and hydranth. Caulus covered by
flexible perisarc that extends slightly onto the hydranth

body. Caulus with gastrodermal canals and at aboral
end with many rooting filaments. Hydranth vasiform,
with one oral and one aboral whorl of tentacles. Oral
tentacles 12-14, moniliform with 4-6 nematocyst
clusters. Aboral whorl with 16-20 filiform tentacles.
Medusae buds develop in clusters on short blastostyles
originating above the aboral tentacles. Medusae buds
naked. Young medusae resemble adult ones, only
without gonads which start to develop soon after
liberation.

Type Locality: Bahamas.

Remarks: Only a limited number of C. forbesii medusae
became available and not all nematocysts could be
examined appropriately. The observed nematocysts
mostly agreed with observations of Bouillon (1978b),
except that the large spherical haplonemes were seen
here as heterotrichous anisorhizas and not as isorhizas.
The distinction between an isorhiza and anisorhiza is
quite often subject to judgment because in large cap¬
sules the thread may taper only gently.

The hydroid stage has not yet been found outside
the Mediterranean. As the medusa is present in coastal
waters of New Zealand, the polyp almost certainly
occurs here. It can be expected on muddy or sandy
bottoms (cf. Brinckmann-Voss 1970).

Records from New Zealand: Leigh Marine Reserve;
Whangateau Harbour (Barnett 1985).

Other Records (medusa): India; Vietnam; southern
Japan; Tortugas; Florida (Kramp 1968); Mediterranean
(Brinckmann-Voss 1970); Seychelles (Bouillon 1978a);
Papua New Guinea (Bouillon 1978b). Polyp known
only from the Mediterranean.

Corymorpha furcata (Kramp, 1948) (Fig. 61a-b)

Euphysora furcata Kramp 1948:19, pi. 1, figs 7-8; Kramp 1957:
5, pi. i fig. 2; Kramp 1959: 89, fig. 40b; Kramp 1961: 40;
Kramp 1968: 15, fig. 30; O'Sullivan 1982: 20, fig. 7, map
4; ?Pages et al. 1992: 19, fig. 17.

Material Examined:

4 medusae from Dana Stri 3624IX, 28°19.5'S, 176°56 f E, 100 m
of wire, 10.12.1928, damaged.

1 medusa from Dana Stn 3627II, 30°08'S, 176°50'W, 4000 m
of wire, 14.12.1928, 6.5 mm high, well-preserved.

Descripiion: Medusa up to 8 mm high, bell-shaped,
higher than wide, with pointed apex. Stomach cylin¬
drical, two-thirds the length of the bell cavity, with a
broad apical chamber. Gonads encircle manubrium
for almost all its length, with longitudinal depressions

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Fig. 61. Corymorpha furcata, from Dana Shi 3627. a) whole
medusa; scale bar 2 mm. b) cnidophore from end of main
tentacle; scale bar 20 pm.

perradially. Four radial canals and circular canal, all
rather broad. Tentacles hollow. One main tentacle,
longer than three times the bell height, bifurcating
twice at the end. Nematocysts confined to ends, carried
on cnidophores with long stalks. One stenotele per
cnidophore. Remaining tentacles shorter, conical.
Tentacle opposite main tentacle longer than other pair.
Polyp stage unknown.

Type Locality: South of Newfoundland Bank.

Remarks: The twice-bifurcating end of the main tentacle
makes Corymorpha furcata easy to recognise. Close
microscopic examination shows the terminal swell¬
ings to be a mass of stenoteles, each carried singly on

a cnidophore (Fig. 61b). Gonads were seen in only a
few specimens. In the animal figured (Fig. 61a), the
gonads were very advanced and encircled the manu¬
brium. Perradial, longitudinal depressions indicated
a division of the gonads. Pages et nl. (1992) described
and figured the gonads as perradial with interradial
interruptions. This is not in accordance with Kramp's
description (1957) and figures (1959, 1968).

Records from New Zealand: Kermadec Islands (Kramp
1965).

Distribution: Widely distributed in the Indian Ocean
and the western Pacific; Chile; Atlantic Ocean between
40°N and 40°S (Kramp 1968).

Corymorpha intermedia n.sp. (Fig- 62a-b)

Material Examined:

4 live medusae collected near Ti Point, Leigh, 8.8.1991.

Ca. 110 specimens collected at Greta Point, Wellington,
surface plankton, dates: 15.12.1993, 20.6.1994, 27.6.1994,
11.10.1994 (last date yielded more than 100 animals, many
mature), some young medusae cultivated until mature,
gametes were obtained and the development followed,
embryos apparently went through resting stage, which,
however, did not develop further in vitro.

10 preserved medusae collected by T. Barnett near Goat
Island, 1983-1984, one selected as holotype FI-640, remain¬
ing medusae deposited as paratype P-1067.

Description: Adult medusa up to 2.5 mm high, with
apical process of variable shape and height, may reach
one-third of the total height. Bell shape varies from
almost spherical to higher than wide, jelly thin.
Peduncle absent. Umbrella margin at right angle to
main axis. Apical canal absent. No nematocyst tracks
on exumbrella. Relaxed velum spanning one-third to
half of radius. Manubrium tubular, measuring half to
three-quarters of the subumbrellar height, mouth
margin with nematocysts, sometimes flaring in pre¬
served specimens. With large oil droplet at base of
manubrium. Gonads encircle manubrium for almost
its entire length, leaving only a small part near mouth
free. Four narrow radial canals ending in 1 large and
3 smaller marginal bulbs, all connected by narrow
circular canal. Only largest bulb bears a single tentacle
with around ten annular and one terminal nematocyst
clusters (moniliform tentacle). Ocelli lacking.

Young medusa spherical, 0.4 mm diameter, with¬
out apical process, no apical canal, no oil droplet or
oil sequestered into many small droplets, 1 tentacle
with 3 annular nematocyst clusters and a single ter¬
minal one. Exumbrella with scattered nematocysts that
are lost later. Nematocysts:

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a) larger stenoteles, (6.5-8.5) x(5-6.5) gm.

b) smaller stenoteles, predominantly around mouth,
(9.5-10.5) x (8.5-9.5J pm.

fi g 62. Corymorpha intermedia n. sp. a) mature medusa
from preserved material; scale bar 0.2 mm. b) nematocysts,
lop row: stenotele 1, stenotele 2, desmoneme, same dis¬
charged, anisorhiza, same discharged; bottom row: atrichous
isorhiza, same discharged, microbasic eurytele, same
discharged; scale bar 10 pm.

c) desmonemes, discharged with three coils, (6.5-
8) x (4.5-5.5) Jim.

d) basitrichous anisorhizas with slowly tapering
thread, (10.5-11.5) x (10-10.5) pm.

e) atrichous isorhizas, (3.5-4.5) x (3.5-4) pm.

f) microbasic eury teles, mostly in juvenile only, swell¬
ing of shaft indistinct, (3.5-4.5) x(3.5-4)mm, s = 1.1.

Colour: transparent.

Polyp stage unknown.

Type Locality: Surface plankton of Goat Island, Leigh
Marine Reserve.

Eiymology: The species name refers to the intermediate
morphology of the medusa which places it between
Corymorpha and Euphysa .

Remarks: Corymorpha intermedia n.sp. resembles medu¬
sae of C, nutans M. Sars, 1835, but clearly differs by
lacking an apical (= umbilical) canal and a peduncle
in all stages. The nematocyst types are mostly the same
for both species, except for the isorhiza which is not
present in C. nutans (Russell 1938a).

Corymorpha intermedia had no apical canal in all
examined medusae. The presence of an apical canal
has previously been used in the generic diagnosis to
distinguish it more clearly from medusae of Euphysa.
The only other difference is the lack of an apical process
in Euphysa medusae. The Corymorpha intermedia
medusa is therefore intermediate between the two
genera as defined by earlier authors; its inclusion in
Conjmorpha is somewhat arbitrary and is made with
reference to the presence of the prominent apical
process. Corymorpha and Euphysa , however, differ
considerably in the anatomy of their polyp stages (see
Bouillon 1985a; Petersen 1990). When more infor¬
mation on the life cycle of C. intermedia becomes avail¬
able, it might become necessary to transfer it to another
genus. Although gametes could be obtained during
this study, few eggs developed and then went into an
apparent resting stage which did not develop further
for a period of two months. The absence of a planula
stage and resting gastrula stages are typical for the
Corymorphidae (cf. Rees 1937; Brinckmann-Voss
1970).

The polyp of C. intermedia will most probably be
found in sandy or muddy bottoms as in many other
species of this genus.

Records from New Zealand: Leigh Marine Reserve
(Barnett 1985, as Steenstrupia sp.; this study); Welling¬
ton Harbour (this study).

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Euphysa Forbes, 1848

Synonyms: (all after Petersen (1990)

Hypolytus Murbach, 1899
Heteractis Allman, 1864
Meiorhopalon Salvini-Plawen, 1987
Euphysomma Kramp, 1962

Solitary hydroids with elongated hydranth. With one
oral whorl of usually four to six short, moniliform or
capitate tentacles, and one to three close-set aboral
whorls of four to sixteen moniliform tentacles, all with
chordal gastrodermis. Hydrocaulus with an irregular
whorl of four to sixteen short papillae, each with a
gastrodermal statocyst and situated just under the
hydranth. Hydrocaulus hollow, without gastrodermal
canals. Perisarcal tube long. With asexual reproduction
through budding of polarity-reversed polyps from the
hydranth and through asexual bodies constricted off
from pointed end of hydrocaulus. Gonophores
develop singly or inclusters just above aboral tentacles.
Gonophores either released as free medusae or remain¬
ing fixed as sporosacs. Medusa with evenly rounded
umbrella, without apical canal. One to four tentacles,
unequally developed but all of the same structure,
moniliform or modified moniliform. Manubrium
stout, cylindrical with small round mouth. Gonads
encircle manubrium.

Type Species: Euphysa aurata Forbes, 1848.

Remarks: This diagnosis was derived from Petersen
(1990). The slight modification made concerns the
length of the manubrium, which in the new diagnosis
may be longer than the bell cavity. This emendation is
necessary to accommodate £. problematica, the only
species of this genus known from New Zealand,

Euphysa problematica n.sp. (Fig. 63)

Material Examined:

1 medusa obtained fromT. Barnett, collected February 1980
or 1981, Whangateau Harbour. Material deposited as
holotype H-643.

Description: Medusa up to 1 mm high, umbrella almost
spherical. Jelly thin, slightly thicker at apex. Manu¬
brium cylindrical, as long as or longer than bell cavity,
with a small apical chamber. Mouth simple, circular.
Gonad encircles manubrium completely and is
restricted to the distal part of the manubrium, leaving
proximal two-thirds of the manubrium free. Four
moderately broad radial canals and ring canal present.
Four broad perradial marginal bulbs with 4 identical
tentacles, these rather short. Tentacles with about 10

Fig. 63. Euphysa problematica n.sp., type specimen; scale
bar 0.2 mm.

nematocyst clusters and a larger terminal cluster
Ocelli absent. Nematocysts: stenoteles, desmonemes
and one additional type of uncertain identity.

Type Locality: Whangateau Harbour near Leigh,
surface plankton.

Etymology: The species name refers to the difficulties
of placing it within the existing system of cory-
morphid classification.

Remarks: This medusa can be assigned to the genus
only with difficulty. It bears some slight resemblance
to Euphysora flammea (Linko, 1905) and £. japonica
(Maas, 1909) in having four identical tentacles. But by
having gonads confined to the distal end of the manu¬
brium it is rather different from all other members of
the genus. In the examined specimen, the manubrium
protrudes from the umbrella; this may, however, be a
fixation artefact. Barnett (1985, as Plotocnide sp.) also
gave a photograph of the only other specimen found
so far. In this specimen the manubrium seems not to
reach beyond the velum.

Although only two specimens are known, and these
in a rather poor condition, the morphology of the
gonads make this medusa easily recognisable. Further
information on the life cycle is needed to confirm its
generic identity.

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Records from New Zealand: Whangateau Harbour,
near Leigh (Barnett 1985).

Family TUBULARIIDAE Allman, 1864

Solitary or colonial hydroids, hydranth vasiform with
one to several close-set whorls of indistinctly monili-
form of pseudofiliform oral tentacles with chordal or
parenchymatic gastroderm, and with one whorl of
long, pseudofiliform aboral tentacles with chordal or
parenchymatic gastrodermis; more or less well-
developed parenchymatic cushion under aboral whorl.
Hydrocaulus divided into a distal neck region covered
by thin periderm, and a proximal stem which may be
short and thick with tuber-like aboral processes, or
long, cylindrical, or cone shaped-shaped with basal
disc or stolons, covered by thicker perisarc.

Gonophores develop on blastostyles arranged in a
whorl just over aboral tentacles. Gonophores may be
released as free medusae or remain fixed as sporosacs.
Free medusae with or without five or eight longi¬
tudinal nematocyst tracks on exumbrella. Umbrella
margin straight or oblique, with one perradial, two
opposite, or four perradial equally developed ten¬
tacles. With or without asexual budding from tentacle
bulb.

Remarks: This diagnosis from Petersen (1990) encom¬
passes the genera Bouillonia, Ectopleura, Hybocodon,
Ralplwria, Tubular in and Zyzzyzus . Only Ectopleura and
Hybocodon are at present known from New Zealand,
but more can be expected. Characteristics of New Zea¬
land species:

Ectopleura : Hydroids with high stems, oral tentacles
in one whorl, perisarc originates from collar on
neck region and does not cover whole neck.
Hybocodon : Hydroids with high stems, oral tentacles
in two whorls, perisarc originates just below
hydranth and covers the whole neck region,
medusa bilaterally symmetrical with one marginal
bulb much larger than the others.

Ectopleura L. Agassiz, 1862

Solitary or colonial hydroids, hydranths vasiform with
oral and aboral tentacles. Oral tentacles arranged in
one whorl. Periderm thin, covering pyriform neck
region and secreted from groove around broadest part
of neck. Hydrocaulus with two, rarely up to five,
internal longitudinal gastrodermal ridges. Stolons
filiform, creeping or forming dense net. Gonophores
arise on blastostyles which are dichotomously
branched or not.

Medusa where present with evenly rounded
umbrella with eight meridional nematocyst tracks
issuing in pairs from tentacle bulbs. Two opposite or
four perradial tentacles, moniliform or with abaxial
nematocyst clusters. Manubrium short, at most
reaching bell margin. Medusa in some species reduced
to radially symmetrical eumedusoids or crypto-
medusoid sporosacs with or without symmetrically
arranged distal protuberances.

Type Species: Ectopleura dumortieri (van Beneden, 1844).

Remarks: The above diagnosis was only slightly
modified after Petersen (1990). Contrary to older
definitions it does not rely on the presence of a free
medusa. The major diagnostic characters distin¬
guishing Ectopleura from other members of the family
are the symmetrical gonophore (medusae or sessile
sporosacs) and the origin of the perisarc in the middle
of the neck region.

There are several very similar Ectopleura polyps
known from the Pacific: £. crocea, E. larynx, E. japonica,
E. radiata, £. venusta, and £. marina. All of these species
have been reviewed by Petersen (1990), For correct
identification it is always necessary to examine mature
female gonophores. Another Ectopleura species that
may be found in New Zealand is £. exxonia (Watson,
1978). This species occurs in Bass Strait, Australia and
is characterised by its small size.

Characteristics of species known from New Zealand:
Ectopleura crocea: polyps with sessile gonophores, in
female with eight laterally compressed distal pro¬
cesses.

Ectopleura larynx: polyps with sessile gonophores, in
female with four rounded distal processes.
Ectopleura nndticirrata : polyps with sessile gonophores,
at least in females with a tuft of tentacle-like distal
processes.

Ectopleura sp.: medusa with eight meridional nemato-
cyst tracks, two opposite tentacles with abaxial
nematocyst clusters, broad marginal bulbs, polyp
unknown.

Ectopleura crocea (L. Agassiz, 1862) (Fig. 64a-g)

Titbulnrin crocea L. Agassiz, 1862: pis 23-23a; Torrey 1902:
42, pi. 3, figs 22-23; Brinckmann-Voss 1970: 28, figs 390-
34 (cum syn.).

Titbulnrin niesembrynnthemum Allman, 1871: 418, figs 83-84;

Hirohito 1988: 18, fig. 4, pi. 1, fig. B.

Ectopleurn crocen : Petersen 1990: 174, fig. 27,

Material Examined:

Live colonies collected on underside of raft near Devonport
Wharf, Auckland, 4.1.1994 (infertile), 13.2.1994 (fertile), 8

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actinulae examined in preserved material, material
deposited.

Preserved material from MoNZ, date and locality of collect¬
ion unknown, but from New Zealand.

NZOI collection, Stn Z1060, 17.8.1954, from New Zealand
probably from ship hull.

Ectopleurn crocea from Cumming's Point, entrance to Charles¬
ton Harbour , South Carolina, USA, preserved, collected,
and identified by Dr Dale Calder, ROM, 2.1.1975, up to
2 cm high.

For comparison, neotype of Ectopleurn rnlphi (Bale, 1884),
Catalogue no. 43227, Museum of Victoria, Melbourne,
Australia, up to 10 cm high.

Description: Colonial, tubulariid hydroids arising from
attached ramifying stolons. Height up to 50 mm. Cauh
not branching, thinner basally (up to one-third of distal
end diameter), with occasional annulated stretches
gastrodermis of cauli with 2 longitudinal ridges
Hydranth broadly vasiform with one oral whorl erf
up to 20 filiform tentacles and one aboral whorl of up
to 22 tentacles with nematocysts concentrated on lows’
side. Between hydranth and caulus a distinct spherica
neck region. On thickest part of neck a filmy perisaic
originates; basal half of the neck, which is covered by
the filmy perisarc, shows a striation pattern. Perisaic
of the caulus firm.

Fig. 64. Ectopleura crocea , only b) from life, a) fully mature female polyp; scale bar 5 mm. b) younger polyps with maturing
gonophores; scale bar 2 mm. c) blastostyle with female gonophores showing variation of the distal crests; scale bar 0.5 mm
d) male gonophore, CL; scale bar 0.2 mm. e) female gonophore, same scale as d). f) actinula from female gonophore, same
scale as d). g) discharged nematocysts: stenotele, anisorhiza, basitrichous isorhiza, desmoneme, microbasic eurytele (foreign?)
scale bar 10 |im.

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Gonophores spherical to oblong, with short stems,
born on blastostyles just above aboral tentacles.
Blastostyles of different length, the largest may bear
op to 35 gonophores and may branch once. Gono¬
phores remain sessile as sporosacs with no radial
canals or circular canals but with spadix. Female
gonophores slightly larger than male ones, with 6-8
laterally compressed processes around distal opening,
size of these crests very variable. Female gonophores
contain 2-6 eggs and up to 2 developing actinulae.
Spadix or tentacles of actinula may protrude from
distal opening of gonophore. Actinula with 8 slightly
capitate aboral tentacles an no rudiments of oral
tentacles. Male gonophores without apical crests,
•ematocysts:

a) larger stenoteles, (6.5-7) x (5-5.5) pm.

b) smaller stenoteles, (5-6) x (3.5-4.5) pm.

c,i heterotrichous anisorhizas, (8-9.5) x (7-9.5) pm.

d) basitrichous isorhizas,(9-9.5) x (3-4) pm.

e) desmonemes, (5-5.5) x (3-3.5) pm.
microbasic euryteles, rare, ca. 9 x 4 pm, s = 0.7.

Measurements: stem diameter basally 0.2 mm, distally
'.6 nun; female gonophore 0.5 x 0.6 mm; male gono¬
phore 0.4 x 0.5 mm.

Type Locality: Boston Harbour, Massachusetts, USA.

Remarks: The morphology of the material referred here
£. crocea accords well with other descriptions of this
species. The only major difference is the presence of
microbasic euryteles in the New Zealand material.
Although the nematocysts had to be examined in pre¬
served material, euryteles were found in both samples
from Devonport taken at different dates. Only a few
discharged nematocysts were found each time. The
undischarged isorhiza and eurytele are very difficult
to distinguish. Euryteles have not so far been reported
for E. crocea (Weill 1934; Bouillon 1985a; Zamponi &
Arca-Tellecha 1988; Petersen 1990), but these sources
do not agree on what types are present. Inasmuch as
the presence of some nematocyst types is easily over¬
looked, especially when there are many as in tubu-
lariids, and also considering that the euryteles of the
>ew Zealand material may have been contaminated,
the examined material is referred to £. crocea. If future
comparison of nematocysts will confirm any dis¬
tinctions, the species from New Zealand may be new.
The presence of microbasic euryteles, however, can
be expected from the cladogram given by Petersen
1990).

The only minor difference from the other des¬
criptions was the occasional occurrence of branching
blastostyles, but in material from the USA some are
also present. Regularly branching blastostyles have
been described for £. ralphi (Bale 1884), a very similar

species from Australia and South Africa (cf. Millard
1975, as T ubularia warreni; Watson 1980; Petersen 1990).
The neotype of Ectopleura ralphi was examined for this
study and it was found that the blastostyles are also
only occasionally branched. The actinulae of £. ralphi ,
however, are reported to have rudiments of oral
tentacles which are absent from £. crocea. Ectopleura
ralphi also seems to form much taller polyps. Until
further, preferably also molecular, analysis has been
made it seems advisable to keep both species separate.

As can be seen from Figure 64c, the distal crests of
the female gonophore of £. crocea may vary greatly in
size and shape. This was also observed by Torrey
(1902) and can make identifications difficult
Ectopleura crocea is distributed by ships, so a wide
distribution can be expected.

Records from New Zealand: Devonport, Auckland.

Other Records: Pacific and Atlantic coast of USA;
Europe; Mediterranean; Japan (Petersen 1990).

Ectopleura larynx (Ellis & Solander, 1786)

(Fig. 65a-b)

Tubiilarin Innjnx Ellis & Solander, 1786: 31; Broch 1916: 27;
Ralph 1953: 68, fig. 12; Brinckmann-Voss 1970: 31 (cum
syn.); Miller 1973: figs A & E, not B; Millard 1975: 35, fig.
15H-J; Werner 1984: fig. 106; Cornelius et nl. 1990: 116,
fig. 4.5

[Not Acharndria larynx Wright, 1863 = Ectopleura wrighti
Petersen, 1979]

Fig. 65. Ectopleura larynx, redrawn from Ralph (1953) with
kind permission of the publisher, a) distal part of polyp;
scale bar 1 mm. b) female gonophore containing a planula
and an actinula larva; scale bar 0.2 mm

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Tubulnhn attenoides Coughtrey, 1876: 302.

Ectopleura larynx. Petersen 1990: 170.

Material Examined:

No material from New Zealand seen; for the description
below one sample was examined originating from Nova
Scotia, Canada, ROM B512, det. D. Calder.

Descrifiion: Colonial tubulariid hydroids, up to 50 mm
high, arising from a dense hydrorhiza formed from
ramifying stolons. Cauli normally not branching, but
with apparent branching caused by settling of larvae
on stems of older polyps. Perisarc of stem firm, with
some irregular annulations which may be more or less
pronounced. Gastrodermis of hydrocaulus with 2-4
longitudinal ridges. Hydranth vasiform with long
hypostome; with one oral whorl of up to 20 filiform
tentacles adnate to hypostome and forming longi¬
tudinal ridges over distal half of hypostome, and with
one aboral whorl of 20-25 long filiform tentacles. Neck
region of hydranth with fine longitudinal striations.
Neck region only covered in lower half by filmy
perisarc secreted from a groove.

Gonophores born on blastostyles that may be
branched or not. Blastostyles arising distal to aboral
whorl of tentacles, as long as tentacles, up to 12 per
hydranth with up to 12 gonophores per blastostyle.
Gonophores remain fixed as sporosacs, up to 0.6 mm
long, oval, without radial canals. Older female
gonophores with 4 closely set distal processes varying
in shape from rounded tubercles to short finger-like
appendages. Male gonophores mostly lack distal
processes.

Additional Information From Lotrature: The actinula
has an aboral whorl of 6-3 tentacles with swollen tips,
oral tentacles may be absent or present as three to five
rudiments (Petersen 1990). Nematocysts (after
Bouillon 1985a): stenoteles, anisorhizas, basitrichous
isorhizas, desmonemes.

Type Locality: British Isles.

Remarks: No New Zealand material could be examined
for this study, but the drawings of Ralph (1953) rather
unambiguously show the specific characters of E.
larynx, a very widespread species distributed by ship¬
ping. Many authors have confused Ectopleura larynx
and E. crocea (Petersen 1990). The identifications are
reliable only where information on the female gono-
phore is provided.

Records from New Zealand: Otago Harbour; Welling¬
ton Harbour (Ralph 1953).

Other Records: Atlantic coast of Europe from White

Sea to Portugal; Iceland; east coast of North America
from Newfoundland to southern New England; Sea
of Okhotsk near southern Sakhalin and the Kuriles
Pacific coast of Washington and British Columbia
(Petersen 1990).

Ectopleura multicirrata n.sp. (Fig. 66a-d)

Material Examined:

Few fertile colonies growing on live Perm canaliculus
(Bivalvia), collected 30-3.1994, 10 m, Steeple Rock beacon,
Wellington Harbour. One colony cultivated for 4 weeks
colonies thriving on Artemia , many new polyps and
gonophores formed, material fixed 30.3.94 and deposited
as holotype H-763.

Description: Colonial tubulariid hy droids, up to 15 mm
high, arising from thick, attached, ramifying stolons
that penetrate shell substratum (into holes made by
boring organisms). Erect part of polyps composed of
caulus, neck region, and hydranth. Cauli unbranched
occasionally annulated or corrugated. Gastrodermis
of cauli with 2 opposite longitudinal ridges. Neck
region forming a spherical or blunt cone. Perisarc
originates from widest part of neck region. Hydranth
vasiform with one oral and one aboral whorl of filiform
tentacles. Oral tentacles 10-14 in number, short and
stubby, not tapering, often 2-3 fused together basallv
basal portions adnate to hydranth body. Aboral
tentacles 16-20 in number, up to 2 mm long, tapering,
oval in cross section, nematocysts concentrated in
narrower sides (abaxial and adaxial). Bases of aboral
tentacles not continued as ridges over hydrant body
Gonophores arise in clusters on unbranched, short
blastostyles arising just above aboral tentacles, 6 or
more blastostyles. Gonophores remain fixed as
sporosacs. Sporosacs elongate, without radial canals
or circular canal but with conspicuous spadix placed
asymmetrically, with one large egg only. Gonophore--
with a distal tuft of up to 30 tentacle-like processes
These processes bear no nematocysts, they are pro¬
vided with a row of chordoid gastrodermis cells. In
young gonophores the processes are short and stubby
and separated into 4 clusters that fuse later. Some tufts
are not exactly positioned at distal end thus rendering
the gonophore bilaterally symmetrical. Male
gonophores not observed. Nematocysts:

a) stenoteles size class 1, rare, (12-13) x (10.5-11) pm,

b) stenoteles size class 2, (8-9) x (7-8.5) pm.

c) stenoteles size class 3, (7-8) x (5.6-6) pm.

d) stenoteles size class 4, (5-6.5) x (3.5-4.5) pm.

e) desmonemes, (5-5.5) x (3-4) pm.

f) basitrichous anisorhizas, very slowly tapering,
almost like isorhiza, (9.5-11) x (3.5-4) pm.

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@®®C

g) anisorhizas with very fine and short spines in Type Locality: Steeple Rock Beacon, Wellington Har-
spirals, rare, (12-13.5) x (9-9.5) pm. hour, depth 10 m.

Colour: greenish in samples from nature, turns orange Etymology: The species name refers to the distal tuft
on Artemia diet. of tentacle-like processes of the gonophores.

Fig. 66. Ectopleura multicirrata n. sp., from life, a) total view of animal with gonophores, at right cross sections of caulus at
corresponding heights; scale bar 1mm. b) neck region with origin of perisarc (dotted), CL; scale bar 0.1 mm. c) nematocysts,
horn left: four stenoteles, desmoneme, discharged desmoneme, basitrichous anisorhiza, same discharged, holotrichous
anisorhiza, same discharged; scale bar 10 pm. d) blastostyle with female gonophores of various stages of development; note
the characteristic tentacle-like structures at the distal end of the gonophores, CL; scale bar 0.2 mm.

Ill

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Remarks: The tuft of up to 30 tentacle-like processes of
the gonophore is unique and not known for any
otherspecies (cf. Petersen 1990). The animals are rather
easy to cultivate in running sea water and feed well
on Arternia nauplii.

Records from New Zealand: Known only from the type
locality, Wellington Harbour.

Ectopleura sp. (Fig. 67)

Material Examined:

Several medusae collected by T. Barnett, Leigh Marine
Reserve, date unknown (1983-1984).

Description: Medusa up to 1.6 mm high and 1.4 mm
wide. Umbrella bell-shaped, narrowing at base. Jelly
moderately thick, about twice as thick apically as
laterally, without clear apical process. Exumbrella
with 4 pairs of longitudinal nematocyst bands origi¬

nating from the sides of the marginal bulb epidermis
Manubrium tubular, about two-thirds as long as bell
cavity, with large apical chamber in the form of a
pointed cone. Gonads encircle the manubrium, leaving
only proximal and mouth parts free. With 4 radial
canals and a circular canal, all rather narrow; 4 broad
marginal bulbs, all the same size, oval in shape
nearly as broad as space between bulbs. Two oppo¬
site perradial tentacles only. Tentacles with one larger
terminal nematocyst cluster and about 6 abaxial
nematocyst clusters distributed along the whole
tentacle. Polyp stage unknown.

Remarks: This medusa resembles Ectopleura minerva
Mayer, 1900b, described originally from Florida. Other
Ectopleura polyps are known that also produce
medusae with two tentacles: E. pacifica, E. larynx
(Wright, 1863) and E. mayeri (cf. Brinckmann-Voss
1970; Petersen 1990). Ectopleura medusae from Leigh
differed from other descriptions of E. minerva in the
absence of a an apical process and the presence of
rather broad marginal bulbs instead of small ones. All
Ectopleura medusae are quite simple and species
distinctions should rather rely on the more complex
polyp stage. Some authors (e.g., Mayer 1910; Hirohito
1988) allocated the medusae E. minerva (type locality
Florida) to the hydroid E. pacifica Thornely, 1900, a
species known from New Britain (Papua New Guinea),
This was based on the unreleased medusa. As stated
by Petersen (1990) this allocation cannot be main¬
tained because there are more Ectopleura medusae
known with two tentacles only. It may well be that
the Ectopleura medusa from Leigh belongs in fact to E*
pacifica, while E. minerva more probably belongs to the
hy droid E. mayeri Petersen, 1990 from Bermuda. For
all species, rearing experiments to elucidate the life
cycle must be made before any 2-tentacled Ectopleura
medusa can be allocated to a certain species. The pre¬
sent material is therefore not assigned a specific name
Also other records of E. minerva from the Pacific should
be seen in this light.

Records from New Zealand: Leigh Marine Reserve
(Barnett 1985, as E. minerva),

Hybocodon L. Agassiz, 1862

Solitary tubulariid hydroids with long, perisarc-
covered caulus and a neck region below hydranth.
Hydrorhiza irregularly branched, slightly swollen and
often embedded in sponges. Perisarc thin, much
inflated around neck region, secreted from groove
between hydranth and neck. Hydrocaulus tubular
with open centre, with eight or more longitudina

112

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gastrodermal ridges. Hydranth vasiform, with two
closely set whorls of oral tentacles and one whorl of
aboral tentacles. Gonophores arise from branching
blastostyles distal to aboral whorl of tentacles and are
released as free medusae. Medusa with bilaterally
■ymmetrical umbrella with oblique margin. Ex¬
umbrella with or without five longitudinal nemato-
cyst tracks. Manubrium cylindrical, on short peduncle,
not extending beyond umbrella margin. With one
shorter, two medium-sized and one longer radial
canals. One broad, bean-shaped marginal bulb with
long moniliform tentacle at end of longest radial canal,
and three equally developed rudimentary bulbs
(modified after Petersen 1990).

Type Species: Hybocodon prolifer L. Agassiz, 1962.

Remarks: Only H. prolifer is known from New Zealand.
Other species from similar southern latitudes are H.
unions and H. cnjptus (cf. Millard 1975; Watson 1984).

Hybocodon prolifer L. Agassiz, 1862 (Fig. 68a-e)

Hybocodon prolifer L. Agassiz, 1862: 243, pi. 23a, figs 10-11,
pi. 25, fig. 19; Mayer 1910: 38, pi. 2, fig. 1, pi. 3, fig. 3,
text-fig. 10; Uchida 1927:192, fig. 29; Fraser 1937: 54,pi.
2, fig. 46; Fraser 1944: 106, pi. 18, fig. 78; Ralph 1953: 70,
figs 11 & 22; Russell 1953: 79, pi. 3, figs 3-4, text-fig. 34;
Kramp 1959: 86, fig. 33; Kramp 1961: 43 (cum si/;?.);
Kramp 1968:12, fig. 19; Arai & Brinckmann-Voss 1980;
10, fig. 4; Roper et nl 1983: table 2; Petersen 1990: 192,
fig. 37; Bouillon 1995: 224.

Fig. 68. Hybocodon prolifer, a) whole polyp; scale bar 5 mm. b) neck region and hydranth with blastostyles; scale bar 1 mm.
c) blastostyle with medusae buds, CL; scale bar 0.5 mm. d) Adult medusa from life; scale bar 0.5 mm. e) nematocysts, from
left: two sizes of stenoteles, basitrichous haploneme, desmoneme, microbasic eurytele, microbasic mastigophore occurring
in polyp only; scale bar 10 Jim.

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Material Examined:

3 polyps collected by P. Ralph, tube No. 235, held at MoNZ,
date and origin not given, but from New Zealand, with
remains of sponge on hydrorhiza, with medusae buds.
Numerous polyps collected live 30.3.1994, Steeple Rock
beacon (Wellington Harbour, near Seatoun), 2 m, on
orange sponge, infertile, cultivated for more than 1 month,
formed blastostyles but no medusae buds.

A few polyps collected live west of TeRaekaihau, April 1994,
2 m, infertile, cultivated for several months.

~10 preserved medusae from Leigh Marine Reserve,
collected by T. Barnett 1983-1984.

~30 live medusae collected near Seatoun 14.5.94, 6.6.94,
28.9.94, 4 of them fully mature, material deposited.

DESCRipriON:

Polyp stage: Solitary tubulariid hydroids, height up
to 60 mm, divided into hydranth, neck region, caulus,
and hydrorhiza. Hydrorhiza formed by branching
stolons embedded in sponges. Caulus tapering proxi-
mally to one-third of distal diameter, with firm peri-
sarc. Gastrodermis with central lumen and several
longitudinal ridges. Neck region simple, not swollen,
surrounded by loose filmy perisarc with wrinkles.
Perisarc originates in groove between neck and
hydranth, turning firm after reaching caulus.
Hydranth pear-shaped, diameter with tentacles
10 mm, with up to 31 longer aboral filiform tentacles
in one whorl, oval in cross section, bases of tentacles
continue as ridges over basal part of hydranth body.
With up to 50 shorter oral filiform tentacles in 2 closely-
set whorls. Oral tentacles continue basally as ridges
on hydranth body. Hypostome rounded. Between oral
and aboral tentacles 8 longer and additional shorter
blastostyles. Blastostyles branching and bearing naked
gonophores that are liberated as free medusae.
Nematocysts:

a) stenoteles, (7-11.5) x (5.5-10) pm.

b) heterotrichous anisorhizas, spines visible for more
than 5 capsule lengths, (11-17) x (10-18) pm.

c) desmonemes, discharged with 3 coils, (5-6) x (3.5-
4) pm.

d) microbasic euryteles with very faint distal swelling
of shaft, some like mastigophores, (9.5-10.5) x (3.5-
4.5) pm, s - 0.8.

e) microbasic mastigophores, rare, (5.5-6.5)x (3.5-
4) pm.

Colour: pink.

Mature medusa: Umbrella bell-shaped, 2-4 mm in
height. Base of bell oblique. Jelly moderately thick,
sometimes with apical depression. Jelly on side of ten¬
tacle thicker. Gastric peduncle present. Dilated velum
spanning one-third of radius. Exumbrella with 5 longi¬
tudinal bands of nematocysts (stenoteles) issuing from
marginal bulbs, tentacular bulb issuing 2 bands.
Nematocyst bands extend almost to apex. Manubrium

simple, cylindrical, reaching velum but not beyond.
Mouth surrounded by narrow ring of nematocysts.
Gonads surrounding stomach, leaving peduncle,
upper portion and distal portion, of manubrium free.
With one short, 2 opposite medium-length and one
long radial canal ending in circular canal, all canals
thin. Longest radial canal ends in a broad, bipartite
bulb with one tentacle. Other marginal bulbs very
rudimentary. Tentacle tapering, with one row of
adaxial nematocyst clasps. Ocelli absent. Nematocysts:

a) larger stenoteles, (8.5-10.5) x (7 8.5) pm.

b) smaller stenoteles, (5.5-7) x (4.5-5) pm.

c) heterotrichous anisorhizas, (8-11.5) x (8-11) pm.

d) microbasic euryteles with indistinct swelling of
shaft, (7-8.5) x (2.5-3) pm.

Colour: bulb and upper end of stomach intensively
red.

Type Locality: Rock ledge off Nahant, north of Boston,
Massachusetts, USA.

Remarks: According to Russell (1953), the peduncle
may be absent, there are not always five nematocyst
bands on the exumbrella, and there may be more than
one tentacle on the large bulb. In younger stages there
may be medusae buds on the tentacular bulb.

Hybocodon prolifer is mostly described as a solitary
species. During the present study, however, some rare
branching of cauli was observed under cultivation.
Despite being solitary, quite a number of polyps can
be found together. The living material examined for
this study did not form medusa buds, only incipient
blastostyles, but the polyp is distinct enough to allow
generic identification even in the absence of medusae
buds. Identification is further aided by the presence
of mature medusae at the same site. The preserved
sample available had medusae buds with meridional
nematocyst tracks and are thus certainly likeH. prolifer.
Medusae buds on medusae were never seen at any
stage. This may indicate a difference from the Atlantic
population. Identification of the anisorhiza is some¬
what uncertain as the thread tapered only very gently,
thus resembling isorhizas. This ambiguity seems to
be a general problem with large haplonemes found in
the Tubulariidae. Haplonemes have not so far been
reproted for H. prolifer , but are known for H. imicus
(Millard 1975; Bouillon 1985a). At present, the obser¬
ved differences (medusa-budding absent, presence of
haploneme) are insufficient to warrant the creation of
a new species.

Records from New Zealand: Polyp stage: Antipdoes
Islands (Ralph 1953); Wellington Harbour and south
coast (this study). Medusa: Avon-Heathcote estuary,
Christchurch (Roper et nl 1983); Leigh Marine Reserve

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(Barnett 1985); Cape Egmont (Bouillon 1995); Welling¬
ton Harbour (this study).

Other Records: Newfoundland to Chesapeake Bay;
West Greenland; Iceland; NW Europe from English
Channel to Beren Island; Bering Sea; Puget Sound: Sea
of Okhotsk; Alaska; Hokkaido to Kamchatka (Arai &
Brinckmann- Voss 1980; Petersen 1990; and Dale Calder
pers. comm.).

Family BOEROMEDUSIDAE Bouillon, 1995

Medusae of the order Capitata, with apical projection,
cylindrical manubrium bearing gonads as four large
perradial pouches hanging freely in the bell cavity.
Four radial canals and circular canal. Four marginal
bulbs, and four simple tentacles with many nematocyst
clusters including a terminal cluster.

Remarks: This family was proposed by Bouillon (1995)
to accommodate the rather unusual Boeromedusa
auricogonia.

Boeromedusa Bouillon, 1995
Diagnosis as for the family.

Type Species: Boeromedusa auricogonia Bouillon, 1995.
Remarks: Only one species is known in this genus.

Boeromedusa auricogonia Bouillon, 1995 (Fig. 69)
Boeromedusa auricogonia Bouillon 1995: 231, figs 3b, 7.
Material Examined:

Holotype H-620, NZOI Stn N433,41°46.1'S, 171°25.9 r E (Cape
Fouiwind), 0-25 m, 30.1.1975.

Description: Medusa 7 mm high and 4 mm wide,
umbrella bell-shaped with widest diameter in the
upper half, with shallow rounded apical process. Jelly
thick, increasing in thickness from margin towards top.
Roof of subumbrella with 4 interradial, solid, pyra¬
midal projections into the mesogloea. Manubrium
Darrel-shaped, length about half of of bell cavity,
circular in cross-section; mouth open, circular with an
inconspicuous ring of nematocysts. Four gonads in
perradial position, originating from base of manu¬
brium; gonads in the form of flattened sac-like pro¬
cesses hanging freely into the subumbrellar cavity.
Each gonad is bent slightly adaxially, longer than wide,

Fig. 69. Boeromedusa auricogonia, holotype; scale bar 1mm.

with distal ends lobed, middle lobe of the 3 lobes
longer than lateral ones. Four radial canals and circular
canal present, all narrow and smooth. Radial canals
widening slightly on entering circular canal. With 4
conical perradial tentacle bulbs that turn without
transition into short tentacles. Tentacles covered by
numerous nematocyst clusters and ending in a large
ovoid terminal cluster. No ocelli observed. Polyp stage
unknown.

Nematocysts (after Bouillon 1995): stenoteles, desmo-
nemes, microbasic euryteles.

Type Locality: Cape Fouiwind, South Island, New
Zealand.

Remarks: Boeromedusa auricogonia is rather unique and
at present difficult to accommodate within the system.
Bouillon's (1995) use of a separate family is thus
followed here. Knowledge of the polyp stage will, it is
hoped, clarify its uncertain systematic position. Re¬
examination of the type species showed only a few

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differences from Bouillon's (1995) description. The
subumbrellar projections into the apical mesogloea
were found to be solid. This was verified by probing
them with a blunt needle. Bouillon (1995) described
them as pockets of the subumbrella. Almost all
medusae in the collection containing B . auricogonia had
lost their ocelli owing to the fixative used. The absence
of ocelli in B. auricogonia is therefore not certain. The
mouth in the only specimen is widely dilated, forming
a circular opening. It may be that the shape of the
mouth differs in live animals.

Records from New Zealand: Known only from a single
medusa taken off Cape Foulwind near Westport
(Bouillon 1995).

Other Records: Not known outside of New Zealand.

Family MARGELOPSIDAE Uchida, 1927

Solitary, pelagic hydroids, adult with filiform tentacles
only. One or more oral whorls of tentacles and aboral
tentacles which are either in several whorls or
dispersed. Occasionally aboral region of hydranth
modified for floating. Gonophores develop between
oral and aboral, or between aboral tentacles and are
liberated as free medusae.

Medusae with no apical projection, no exumbrellar
tracts of nematocysts. Mouth simple, circular, no lips,
no oral tentacles. Gonads encircle manubrium com¬
pletely. Four radial canals. Tentacles solid, generally
moniliform, distributed either in perradial groups on
equal height, or in pairs at different levels of exum¬
brella. No ocelli present. Eggs may develop into
actinula on manubrium ( Climacocodon , Margelopsis).

Remarks: The above diagnosis was slightly modified
after Bouillon (1985a) by allowing more than one
whorl of oral tentacles in the polyp, a condition seen
in Pelagohydra . Only the genus Pelagohydra is known
from New Zealand.

Pelagohydra Dendy, 1902

Margelopsid hydroid with aboral half of hydranth
transformed into floating body, without caulus. Adult
medusa unknown, young ones with four groups of
five to seven marginal tentacles. Each group composed
at least of two long abaxial and three short adaxial
tentacles. Monotypic genus (Bouillon 1985a).

Type Species: Pelagohydra mirabilis Dendy, 1902.

Pelagohydra mirabilis Dendy, 1902 (Fig. 70a-d

Pelagohydra mirabilis Dendy 1902: 1, pis 1-2; Mayer 191(h
83, fig. 40a-b; Percival 1938: 439; Ralph 1953: 66, figs 7
23; Kramp 1961: 50; Pilgrim 1967a: 439, figs 1-6; Pilgrim
1967b: 491; Kramp 1968: 18, fig. 40; Rees & Ralph 1970:
11, pis 1-2, fig. 4; Bouillon 1974: 143; Roper et al. 19fl3
table 2; Wemer 1984: 178, fig. 98.

Material Examined:

Polyp and liberated medusae collected and described by R.
Pilgrim (1967), 22.4.1966, from Sumner Beach, held at
MCC.

Description:

Polyp stage: Solitary hydroids freely floating in the
sea, up to 35 mm long. The body of the hydranth is
divided into a larger oval part (float) and a smaller
tubular oral part (proboscis). The float bears up to 150
scattered, tapering tentacles, length up to 7 mm. The
oral part of the hydranth is provided with up to Si
tentacles scattered over the distal three-quarters of its
length. These oral tentacles are shorter than the aboral
ones and slightly capitate; towards the mouth they
originate very obliquely so that their basal part
becomes adnate to the proboscis wall. Along the mouth
rim are some very short, differently coloured ten¬
tacles. The float has a complicated internal anatom)
consisting in an intricate structure of mesogloea!
lamellae and gastrodermal chambers (for details see
Dendy 1902; Rees & Ralph 1970).

Gonophores develop on branched blastostyle-
which are dispersed between the aboral tentacles, up
to 300 per animal. The blastostyles may bear up to 5
gonophores which develop into free medusae.
Nematocysts:

a) stenoteles, (11-13) x (9-10) pm.

b) desmonemes, (6-7) x (4-4.5) pm.

Young medusa:With a bell-shaped umbrella (1.4 mm,
preserved), jelly rather thick (0.2 mm), apex only
slightly thicker. Exumbrella with many scattered
stenoteles and haplonemes. Manubrium cylindrical^
reaching velum or projecting beyond it, manubrium
base quadratic with apical canal, mouth simple. No
gonads visible. With 4 radial canals and a circular
canal. Four large perradial marginal bulbs; these
extend from velum around umbrella margin onto
exumbrella. Each tentacle bulb has 6-7 slightly
capitate, solid tentacles in a special arrangement: the
most abaxial pair points sideways, the next pair
projects downwards and is followed adaxially by a
single median tentacle which also projects down¬
wards and then 1-2 small tentacles projecting
adaxially. Nematocysts:
a) stenoteles, (10-15) x (8-12) pm.

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cA(T

b) desmonemes, 6x4 \im.

c) heteronemes, (7-8.5) x (2.5-3) \xm.

d) haplonemes (basitrichous isorhizas?), (6-7) x (4-
5.5) Mm.

Adult medusa: unknown.

Type Locality: Sumner Beach, Christchurch, New Zea¬
land.

Remarks: Pelagohydrn mirabilis is a rarely reported and
unique species. The rarity may, however, be due to
the paucity of plankton studies in New Zealand. Roper
ct al. (1983) found several polyps and medusae in the
Avon-Heathcote estuary in Christchurch. It may well
be that the species is more frequent than previously
thought. The distribution of P. mirabilis is restricted
to N ew Zealand and all animals found so far have

been found near Christchurch. This very localised dis¬
tribution is rather astonishing considering that the
polyp is freely floating. It would be very interesting
to know the complete life cycle and to have more
ecological data.

The peculiar structure of Pelagohydrn has attracted
much interpretation and speculation (e.g., Garstang
1946). Rees and Ralph (1970) re-examined the anatomy
and convincingly homologised the float region to the
aboral end of a tubulariid hydranth and the proboscis
to the oral region. They also made clear that Pelago-
hydra has no caulus like sessile forms, and offered a
model on how the float evolved.

Records from New Zealand: Christchurch area; Porto-
bello (W. Vervoort, pers. comm.).

Other Records: Not known outside of New Zealand.

Fig. 70. Pelagohydra mirabilis , from preserved material a) floating polyp stage (mouth downwards) with some medusae
buds; scale bar 2 mm. b) young medusa; scale bar 0.5 mm. c) oral view of tentacle bulb of young medusa, the adaxial side
faces upwards; rc radial canal, te tentacle; scale bar 0.1 mm. d) nematocysts of medusa: stenotele, heteroneme (eurytele?),
desmoneme, basitrichous isorhiza (?); the polyp stage has only stenoteles and desmonemes; scale bar 10 Jim.

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Family CANDELABRIDAE de Blainville, 1830

Solitary hydroids; hydranth elongated, cylindrical
with thickened mesolamella and gastrodermal villi;
numerous scattered, hollow capitate tentacles, simple
or compound. Hydrocaulus short, stout, with tubular
or root-like adhesive processes, with or without peri-
sarc; gonophores develop directly on hydranth or on
coryniform blastostyles in budding zone under body
tentacles.

Remarks: All known species of the family reproduce
by fixed sporosacs. Only the genus Candelabrum is
known from New Zealand. (For a systematic review
of the genus see Segonzac & Vervoort 1995.)

Candelabrum de Blainville, 1830

Hydroids with long, cylindrical hydranth with
numerous densely packed, separate tentacles; hydro¬
caulus plate- or tuber-like, with adhesive processes
that end in discs covered by firm, lamellar perisarc.
Gonophores borne on coryniform blastostyles
developed from aboral part of hydranth.

Type Species: Candelabrum cocksii (Vigurs, 1849),

Remarks: Only one species of Candelabrum is known
from New Zealand.

Candelabrum sp. (Fig. 71a-b)

Material Examined:

1 specimen collected from west side of Te Raekaihau,
11.4.1994, 1 m, growing on stone underneath layer of
encrusting coralline algae, cultivated on glass slide for 8
months in running sea water, fed on Artemin nauplii.
Animal grew from 7 to 25 mm. No gonophores formed.
Preserved animal deposited.

Description: Solitary hydroid, when fully extended
reaching more than 25 mm in height. Shape very
variable, mostly club-shaped with broad end as base.
Body of hydranth except most basal part covered by
a large number of capitate hollow tentacles. Tentacles
especially dense around oral region, becoming gradu¬
ally shorter proximally. Basis of hydranth with up to
3 short, stout processes that anchor polyp to sub¬
stratum. Near base often up to 4 long, tentacle-like
processes that end in a sucker-like terminal swelling.
These tentacles can attach to the substratum and then
transform into the stout anchoring processes or are
again reduced. No perisarc discs were observed at the
end of anchoring processes. No gonophores formed.

Fig. 71, Candelabrum sp. from life, a) young polyp; scale
bar 0.5 mm. b) nematocysts: microbasic eurytele, same dis¬
charged, stenotele, large desmoneme, small desmoneme;
scale bar 10 jim.

Polyp has limited mobility. Nematocysts:

a) stenoteles, (10-12) x (8-10) |im.

b) smaller desmonemes, discharged with 4 coils, (7-
8.5) x (4.5-5.5) pm.

c) larger desmonemes, (10-11.5) x (8-9) pm.

d) microbasic euryteles, (14-17) x (5.5-6.5) pm, s =

1 . 1 .

Remarks: This Candelabrum cannot be identified to
species level, owing to absence of information on gono¬
phores. There are a number of Candelabrum species
known from the southern hemisphere (cf. Briggs 1939;
Manton 1940; Millard 1971, Millard 1975; Segonzac &
Vervoort 1995) and the gonophores are crucial in
separating the species. The only specimen found in

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this study may not have been fully grown. It also
lacked perisarcal discs at the attachment sites, which
may be a culture artifact. It was able to change position
and in one month moved about 1 cm.

Records from New Zealand: East side of Te Raekaihau
this study).

Family CORYNIDAE Johnston, 1836

Capitate hydroids forming stolonal or erect, branching
colonies, arising from attached, ramified stolons.
Perisac firm. Hydranths on cauli, with one whorl of
capitate oral tentacles and often below them more
capitate tentacles in whorls or scattered. There may
be filiform tentacles below the capitate ones. Gono-
phores develop on the polyps, proximal to or among
the tentacles, either sessile sporosacs or are liberated
as free medusae.

Medusae are bell-shaped, with a simple circular
mouth, four radial canals, circular canal, and 2 to 4
hollow marginal tentacles. The gonads encircle the
manubrium completely. Mostly with abaxial ocelli on
the tentacle bulbs.

Remarks: In this memoir the traditional system of
classification (Bouillon 1985a) is used in preference to
that based on phylogenetic analysis by Petersen (1990).
Although the present system is certainly artificial,
Petersen's classification of the Corynidae is difficult
to use and some characters need revisions (see also
discussion in Kubota & Takashima 1992; Pages et al.
1992). The material in the present study shows especi¬
ally the site of medusae budding and the shape of
marginal bulbs are not as simple to use as proposed
by Petersen (1990). Although it is highly desirable to
create a classification basing on phylogeny, in the case
of the Corynidae it seems advisable to wait until, for
example, a RN A sequence comparison confirms Peter¬
sen s cladogram.

Characteristics of New Zealand genera:

Coryne : polyps with many capitate tentacles and ses¬
sile sporosacs.

Diptirena: free medusae with manubrium projecting
beyond umbrella, gonad split into several rings.

> arsia : polyps with many capitate tentacles and pro¬
ducing free medusae with gonads not interrupted.

Conjne Gaertner, 1774

Colonial polyps with capitate tentacles all over their
body, either scattered or in more or less distinct
whorls; sometimes with a whorl of filiform tentacles

beneath the capitate ones. Gonophores arise from the
hydranths and remain fixed as sporosacs (after
Bouillon 1985a).

Type Species: Conjne pusilla Gaertner, 1774.

Remarks: Diagnostic characters of New Zealand
species:

C. pusilla: branching colonies, tentacles not in distinct
whorls, without perisarc funnel beneath hydranth,
without mastigophores and desmonemes.

C. tricycla : branching colonies, tentacles in three
distinct whorls, with perisarc funnel beneath
hydranths, without mastigophores and desmo¬
nemes.

ICoryne sp. 1: stolonal, not branching colonies, ten¬
tacles not in distinct whorls, without perisarc
funnels, with mastigophores and desmonemes.
ICoryne sp. 2: stolonal, not branching colonies, ten¬
tacles in two sometimes three distinct whorls each
containing four tentacles, with basitrichous
isorhiza.

Conjne pusilla Gaertner, 1774 (Fig. 72a-d)

Coryne pusilla Gaertner, 1774: 40, pi. 4, fig. 8; Broch 1916:16;
Stechow 1919: 5, fig. A; Brinckmann-Voss 1970: 51, fig.
57; Morton & Miller 1973: 155, fig. 55; Millard 1975: 51,
figs 19F-G; Boero 1981: 187, fig. 1C-D; Hirohito 1988:
33, fig. lOa-e.

Material Examined:

Several colonies on underside of raft near Devonport Wharf,
Auckland, collected 13.2.1994, female gonophores present.

Description: Hydroid colonies with branching stems,
arising from attached, ramified stolons. Stems up to
20 mm high and with up to 20 hydranths, irregularly
branching in all planes, branching mostly at acute
angle. Perisarc green (preserved material), occasion¬
ally annulated, especially so at origin of side branches.
Perisarc ends below hydranth; no basal calyx.
Hydranths are spindle-shaped, up to 1.2 mm, with
dome-shaped hypostome encircled by 4 capitate ten¬
tacles, tapering only slightly when fully expanded;
below oral tentacles up to 16 capiate tentacles in 3-4
indistinct whorls. No filiform tentacles. Gonophores
spherical, arising on short stems in the upper axils of
the proximal tentacles, occasionally 2 gonophores of
different age per tentacle; mature gonophores slightly
oval (only female seen) remaining fixed as sporosacs,
lacking radial and circular canals. A hollow spadix is
present, surrounded by approximately 25 polygonal
eggs. The mature female gonophore has a terminal
knob without nematocysts. Nematocysts:

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a) larger stenoteles, (21-23) x (13-15) mm,

b) smaller stenoteles, (10-12) x (5.5-6,5) mm.

Remarks: The present material agrees rather well with
other descriptions from the more recent literature,
except that the annulation was not as pronounced as
indicated by Brinckmann- Voss (1970) or Hirohito
(1988). Most probably the extent of annulation is
dependent on the environment.

Fig. 72. Coryne pusilla, from preserved sample, a) colony
form; scale bar 2 mm. b) nematocysts: two sizes of steno¬
teles; scale bar 10 pm. c) hydranth with gonophores, CL;
scale bar 0.5 mm. d) female gonophore arising from the
upper axil of tentacle, CL; scale bar 100 pm.

The presence of C. pusilla in New Zealand was
indicated by Morton and Miller (1973), who gave no
locality. The only sample found in the present study
came from the underside of a raft at Devonport wharf
where it occurred with Pennnria disticha and Ectopleura
crocea. All of these species are distributed by ships.

Records from New Zealand: Devonport, Auckland (this
study).

Other Records: All European coasts; Mediterranean
South Africa; Indian Ocean; Korea; Japan (Brinck-
mann-Voss 1970; Hirohito 1988).

Coryne tricycla n.sp. (Fig. 73a-d)

Coryne vaginata: Ralph 1953: 66, fig. 13.

Material Examined:

Holotype colony H-641, collected 5.10.1994 east of Te
Raekaihau, Wellington, growing on intertidal rock, fertile
female, 45 mm high.

Description: Hydroid colonies arising from attached,
ramified stolons. Stems up to 45 mm high with up to
30 hydranths. With monopodial growth and branching
in all planes. Perisarc soft and elastic, annulated
throughout. At the base of the hydranths the perisarc
dilates to a thick, gelatinous funnel into which
hydranths unable to retract. Hydranths up to 2 mm
long (free part), cylindrical to slightly pear-shaped;
with an oral whorl of 4 short, adnate capitate tentacles,
one median whorl of 4 capitate tentacles in alternate
positions to oral tentacles, and one aboral (proximal)
whorl of 8 capitate tentacles that originate all at the
same level from a slightly thickened region of the
hydranth body; they are disposed in the same plane.
The median whorl is either exactly in the middle of
the body or closer to the oral whorl of tentacles.
Gonophores arise just above the aboral whorl of ten¬
tacles and alternate in position with these; there are
up to 3 gonophores per position, all in various stages
of development. Mature female gonophores are
oblong and remain fixed as sporosacs with up to 100
eggs; a spadix is present but radial or circular canals
are absent. Male gonophores unknown.
Nematocysts:

Only stenoteles of very variable size, (19-36) x (9-
22) pm.

Colour: brown, especially gonophores. Stem diameter
0.20-0.23 mm, gonophores up to 1 mm long, size of
spawned eggs 114 jam (s.d. = 6 jam, n = 6).

Type Locality. Northeast of Te Raekaihau (Lyall Bay,
Wellington), intertidal.

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Etymology: The species name refers to the regular
arrangement of the tentacles in three whorls.

Remarks: At first glance, Coryne tricycla can be mistaken
for Coryne muscoides (Linnaeus, 1761) (of which C.
vaginata is a synonym). Both species have a similar

colony form and the characteristic funnel-shaped
dilatation of the perisarc proximal to the hydranths.
A closer look reveals, however, that they must be
different species, if not different genera. In contrast to
C. muscoides , C. tricycla has a different and unique
arrangement of the tentacles in three well-defined

Fig. 73. Coryne tricyla n.sp., from live holotype. a) distal half of colony; scale bar 2 mm. b) young hydranth with median
tentacle whorl not yet formed; scale bar 0.5 mm. c) intermediate stage with growing gonophores and incipient median
tentacle whorl, same scale as b). d) hydranth with mature gonophores and median tentacles fully developed, same scale as
b. e) nematocysts: stenoteles of various sizes; scale bar 10 |im.

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>®@©

whorls with a constant number of tentacles. The most
proximal whorl with its eight tentacles is especially
unique. Further differences are found in the inability
of the hydranths to retract into the perisarc funnel,
the more oblong gonophores in C. tricycla (cf.
Cornelius etal. 1990) and their position (inC, muscoides
in the axils of the two proximal whorls of tentacles),
the possible occurrence of desmonemes inC. muscoides
(Brinckmann-Voss 1970, but cf. Boero 1981), and the
sequence of development of the tentacles. In the
examined mature colony of C. tricycla, the median
whorl of tentacles was formed last, after the gono¬
phores started their development (Fig. 73b-d). In C.
muscoides, the sequence of tentacular development is
from distal to proximal (Brinckmann-Voss 1970).

The slightly swollen region of the hydranth body
at the level of the aboral whorl of tentacles is some¬
what reminiscent of hydranths of Pennaria, but the
structure of the tentacles, and the cnidome, preclude
any association. The unique morphology of the aboral
tentacle whorl could be used as reason to create a new
genus for this species.

Ralph (1953) reported C. muscoides (as C. vaginata)
from near Wellington. Her figures are compatible with
C. muscoides but not with C. tricycla . However, the
drawings may not be accurate and it seems much more
probable that she had C. tricycla before her because C.
muscoides is known only from Europe. Her record of
C. muscoides in New Zealand should therefore be
treated as a misidentification until confirmation.

Although forming large colonies, C. tricycla is not
so easy to detect as its brown colour perfectly blends
into the surrounding colours of macroalgae.

?Coryne sp. 1 (Fig. 74a-b)

Material Examined:

1 colony growing on stone collected 11.4.1994 west of Te
Raekaihau, 1 m, cultivated for 4 months, after 9 days
gonophores started to grow in axils of tentacles, but
aborted later at very early stage. So me material deposited.
1 colony growing on sponge underneath encrusting bryo-
zoan, collected 15.12.1994, Barretts Reef, Wellington, 10 m.

Description: Hydroid colonies arising from attached,
ramified stolons. Hydranths on unbranched stems.
Stolons and stems covered by firm perisarc without
regular annulations. Hydranths spindle-shaped with
one oral whorl of capitate tentacles and up to 18
capitate tentacles in up to 5 indistinct whorls below.
Oral tentacles somewhat adnate. All tentacles rather
short and stubby in colonies from nature, becoming
gradually shorter and smaller proximally, the most
proximal tentacles with almost no terminal swelling,
but nematocysts confined to tips. No filiform tentacles.

Gonophores most probably arise in the upper axils of
the lowest 2-3 whorls of tentacles, but only initial
stages observed. Nematocysts:

a) larger stenoteles, (39-40) x (21-24) pm.

b) medium size stenoteles, (24.5-28)x (15.5-18) pm.

c) smaller stenoteles, (12.5-15) x (7.5-9.5) pm.

d) mastigophores, shaft only slightly thicker than
thread, (14-16) x (5-6) pm, s ~ 0.8.

e) desmonemes, thread in undischarged capsules
with ropy texture, with 4 coils when discharged,
(10-12) x (5-6.5) pm.

Measurements: hydranths up to 2.4 mm high and
0.4 mm diameter; stems 1-3 pm high.

Remarks: This species is easily distinguished from all
other known corynids known from New Zealand by
its size, the stubby tentacles, and its nematocysts.
Because the nature of the gonophores remains un¬
known, the species was notgivena specific name. With
the presumable gonophores in the upper axils it

Fig. 74. ?Coryne sp. 1, from life, a) polyps shortly after
collection from nature; scale bar 0.5 mm. b) nematocysts,
from top: large, medium-sized and smaller stenotele,
mastigophore, same discharged, desmoneme, same
discharged; scale bar 10 pm.

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resembles Coryne sensu Petersen (1990). It is therefore
provisionally assigned to Coryne, pending more
knowledge of the life cycle. This species is rather
distinct from the majority of corynids as it contains
desmonemes (cf. Petersen 1990). It may therefore also
belong to a completely different capitate group (e.g.,
?Tiaricodon).

Colonies were kept in running sea water on their
original substratum and were rather easy to cultivate.
They did not feed on Artemi a, but voraciously con-
umed large planktonic copepods. The colonies
needed only occasional feeding, yet with even more
food failed to develop gonophores beyond initial
-tages. These initial stages were only bulges of the size
of the tentacular capitae. In culture, the tentacles grew
about twice as long as those seen in nature.

Records from New Zealand: Te Raekaihau, Barretts
Reef (this study).

?Coryne sp. 2 (Fig. 75a-b)

Material Examined

colony growing on unidentified chitinous tube, one colony
growing attached on sand grain, both found 11.11.1994
on surface of sand in 1 m of water at Greta Point, Evans
Bay Wellington collected 2.11.1994; one colony cultivated
in 5 ml plastic petri dish at ambient sea temperature until
3.1,1995, no gonophores formed.

Description: Hydroid colonies arising from ramifying
-tolons. Cauli not branched, up to 1 mm. Stolons and
cauli covered by perisarc. Perisarc of caulus thin,
-lightly funnel-shaped, with adhering silt and detritus.
Hydranth club-shaped, reaching 0.5 mm high, with
nipple-shaped hypostome. With 2, sometimes 3 pre-

Fig. 75. ?Coryne sp. 2, from life, a) hydranth as found on
sand; scale bar 0.2 mm. b) nematocysts: stenotele, basi-
trichous isorhiza, same discharged; scale bar 10 |im.

cise and closely set whorls of capitate tentacles. Four
tentacles per whorl, alternating in position with the
other whorls. Tentacles long, with few chordoid
gastrodermis cells. Nematocysts:

a) larger stenoteles, (17-19) x (12.5-13) pm.

b) smaller stenoteles, (13.5-15) x (9-10) pm.

c) basitrichous isorhizas, (12-15) x (3-4) pm.

Remarks: In the absence of gonophores, no definite
generic name can be given. It is therefore also not
named specifically. The absence of desmonemes and
the tentacles in more than one whorl are in accord
with the Corynidae, and the species was therefore
provisionally referred to Coryne. Although the genus
remains uncertain, the species is most probably new.
The tentacles are in rather exact whorls (2-3), each
containing four tentacles. The arrangement of tentacles
is very regular in all hydranths. Such a configuration,
together with the basitrichous isorhizas make this
species rather distinct. The colonies were found on
the surface of a sand sample that had been standing
submerged for nine days. Most probably the colonies
developed during that time because initially no
hydroids could be detected. Whether sand is the
natural habitat of this species is not certain. The polyps
feed well on Artemia nauplii and the colony grew very
rapidly. Unfortunately no gonophore formation was
observed, even after transfer to running sea water.

Records from New Zealand: Evans Bay, Wellington
(this study).

Dipurena McCrady, 1859

Colonial hydroids with one or several whorls of
capitate tentacles, with or without one whorl of aboral
filiform tentacles. With a button of high gland cells
around mouth. Polyps often associated with sponges.
Gonophores released as free medusae. Gonads of
medusae in two or more rings around manubrium
(after Bouillon 1985a).

Type Species: Dipurena strangulata McCrady 1859.

Remarks: Bouillon (1971) and Pages et al. (1992) gave
a summary of the characteristics of Dipurena medusae
and polyps. Only D. ophiogaster is known from New
Zealand.

lipurena ophiogaster Haeckel, 1879 (Fig. 76a-d)

1 iparena ophiogaster Haeckel, 1879: 25; Uchida 1927: 187,
fig. 27; Rees 1941: 131, fig. 2; Russell 1953: 71, pi. 1, fig.

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5, pi. 2, fig. 4, text figs 25e, 30a-b, 31; Kramp 1959: HC
fig. 22; Kramp 1961: 23 (cunt si/#/.); Kramp 1968: 8, fig
11; Petersen 1990: 212, fig. 44E; Pages et ni 1992: 16.
Slabberia ophiogaster: Mayer 1910: 79, figs 36-37.

Snrsia ophiogaster : Brinckmann-Voss 1970: 59, pi. 3 fig. 4
text-figs 66-71.

Material Examined:

4 (24.11.1993), 10 (9.12.1993), 5 (12.1.1994), 3 (31.1.1994)
(7.3.1994), medusae from Evans bay plankton, most oi
them mature; gametes were spawned and about 3l
planulae obtained. Resulting planulae settled on bottom
of plastic petri dish and metamorphosed into primary
polyps with long cauli. The oldest primary polyps had
one whorl of four capitate tentacles and rudiments of
tentacles in another whorl. They fed only reluctantly and
the colonies deteriorated afterwards.

Description:

Medusa stage: Mature medusa 4-5.5 mm high, bell¬
shaped, 1.5 times higher than wide. Jelly unevenly
thick, becoming gradually thicker from margin
towards top; at apex 3 times as thick as lateral walls
Relaxed velum spanning half to two-fifths of radius.
With a distinct, rounded apical chamber. Manubrium
very long, up to 3 times the bell height, with long and
thin proximal part and a broader distal stomach-
Gonads distributed in 2-4 broad rings encircling
manubrium. With 4 narrow radial canals and ring
canal. Four marginal bulbs, epidermal part with
nematocysts, rather flat, bearing an abaxial, dark-
brown ocellus; gastrodermal chamber egg-shaped
circular canal entering adaxially. With 4 very long and
thin tentacles, length up to 5 times bell height, with a
up to 70 nematocyst clusters and a slightly larger
terminal cluster. Nematocysts:

a) larger stenoteles, (14-15) x (9.5-10.5) |im.

b) smaller stenoteles, (11-12.5) x (8-9) pm.

c) desmonemes, with ropy thread in undischarged
capsules, (11-13) x (5.5-6.5) pm.

Egg size (spawned): 122 pm (n = 5, s.d. = 5.7 pm).

Polyp stage and young medusa (after Rees 1941
Brinckmann-Voss 1970): Colonial hydroids with rarely
branching stems arising from attached, ramifying
stolons. Perisarc covers stolons and stems, without
annulation. Stems (cauli) may reach 2 mm in height
with wide perisarc into which basal part of polyp can
retract (colonies from nature, not cultivated ones)
Hydranths 1-2 mm long, with 10-18 capitate tentacles,
scattered or in indistinct whorls. With 4 aboral filiform
tentacles that may also bear occasional nematocysts

Fig. 76. Dipurena ophiogaster from life, except d). a) mature medusa from Wellington Harbour; scale bar 1 mm. b) side view
(top) and frontal view (below) of marginal bulbs; scale bar 0.3 mm. c) nematocysts of medusa: two stenoteles, desmoneme;
scale bar 10 pm. d) polyp stage with medusae buds, redrawn from Rees (1941) with permission of Cambridge University
Press, no scale given. Note: the polyp stage has not yet been identified from New Zealand waters.

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Filiform tentacles often absent or contracted in colonies
fr om nature, always present in cultures. Medusae buds
develop in groups of up to 4 on short blastostyles
distally to filiform tentacles.

Newly released medusa spherical, approximately
1 mm in size, with evenly thin jelly, with apical canal,
with 8 adradial rows of exumbrellar nematocysts.
Manubrium length one-third of bell cavity, simple
cylindrical.

Nematocysts of polyp (Bouillon 1985a): stenoteles
only.

Type Locality: First described from British Isles and
Ireland.

Remarks: With its very long manubrium, Dipurenn
ophiognster is rather distinct from all other New
Zealand medusae. Adult D. ophiognster are, however,
very difficult to distinguish from D. reesi, a medusa
known from the Mediterranean and Brazil. Brinck-
mann-Voss (1970) gave some help in discriminating
them. Both species are easy to distinguish in their
polyp phase, as D. reesi polyps have only one whorl
of capitate tentacles, making them similar to polyps
erf Cladonemn radiatum (Fig. 80c). Polyps of D. ophio-
gaster, on the other hand, resemble certain other
Dipnrena polyps and also Sarsia japonica, especially
when without medusae buds (Fig. 79a).

Polyps of D. ophiognster have not yet been found in
New Zealand. According to Brinckmann-Voss (1970)
they occur in shaded parts of the rocky littoral 1-3 m
deep on rocks or on barnacles.

Records from New Zealand: Leigh Marine Reserve,
Whangateau Harbour (Barnett 1985), Wellington
Harbour (this study).

Other Records: Northwestern Europe; Mediterranean;
Sri Lanka; Japan; Palau Islands; Papua New Guinea;
Pacific Coast of Mexico and Chile (Kramp 1968).

Sarsia Lesson, 1843

Hydranths similar to Coryne, but with free medusae.
Medusae with undivided gonads (Bouillon 1985a).

Type Species: Sarsia tubulosa (M. Sars, 1835).

Remarks: Characteristics of New Zealand species:

S. exuni a: polyp colonies always branching, without
filiform tentacles.

S. japonica: polyp colonies mostly stolonal, with
filiform tentacles.

Sarsia eximia (Allman, 1859) (Figs 77a-h, 78)

Coryne eximia Allman, 1859:141; Petersen 1990:211, fig. 43 A-
C.

Coryne tenelln Farquhar, 1895: 208, pi. 13, fig. 5 (new syno¬
nym).

Sarsia eximia: Mayer 1910: 57, fig. 20; Hartlaub 1914: 8, figs
1-2; Russell 1938b: 150, figs 8-12; Ralph 1953: 74, fig.
24; Russell 1953: 50, text fig. 17A, 18A-B, plate 2 fig. 3;
Kramp 1959: 79, fig. 15; Kramp 1961: 27; Kramp 1968: 7,
fig. 6; Millard 1975: 52, fig. 20A-D; Brinckmann-Voss
1989: 688, figs 5-6.

Syncoryne tenella: Bale 1924: 228; Ralph 1953: fig. 15.
Syncoryne eximia: Bale 1924: 229; Ralph 1953: 68.

Material Examined:

Type material of Ccnyne tenella Farquhar, 1895, MCC type
AQ3276, type locality Wellington Harbour. Short des¬
cription: erect colonies arising from ramifying hydrorhiza,
fragments up to 12 mm high, branching more distally, up
to 10 terminal branches ending in hydranths. Perisarc
strongly annulated for short, dispersed stretches,
especially at origin of branches; hydranths up to 1.5 mm,
tubular to spindle-shaped body with 4 oral and 15-22
scattered capitate tentacle below them, no basal cup of
perisarc present; only a few young gonophores present,
these in the upper axil of the proximal tentacles, with short
stems. Additional data: gastrodermal cells in oral tentacles
13-15; hydrorhiza diameter 176-216 Jim, stem proximally
110-208 Jim, caulus at hydranth base 104-143 Jim,
hydranth length 900-1480 |im.

Numerous living colonies from different localities in Well¬
ington Harbour, present year-round, very often with
medusa buds, medusae released from several colonies and
cultivated to maturity (spawning); some of the polyp
material deposited.

Many hundreds of adult medusae from Wellington Harbour
(Evans Bay, Seatoun), present year-round; some deposited.
Ca. 10 medusae collected by T. Barnett at Leigh Marine
Reserve in 1983-1984, identified as S. eximia, S. gracilis and
Sarsia sp.

Description:

Polyp stage: Capitate hydroids forming initially
stolonal, later erect, branching colonies arising from
attached, ramifying stolons. Colonies reaching 40 mm
in height, but mostly 10-20 mm. Stems not fascicled,
branching irregularly with an occasional tendency to
unilateral arrangement. Perisarc mainly smooth but
annulated at base of stem, at origin of branches and
sometimes for the entire length of small branches. The
perisarc terminates as a very delicate layer below the
most proximal tentacles.

Hydranths are tubular to spindle-shaped and reach
2 mm in length, normally 1.5 mm; with one whorl of
4-5 oral tentacles; beneath are 15-35 additional ten¬
tacles, scattered or in indistinct whorls; tentacles
capitate and proximal part tapering only slightly. Oral
tentacles at their point of insertion shorter than twice
the diameter of the hydranth. Filiform tentacles absent.

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Fig. 77. Sarsia exitnia from life, a) colony forms; scale bar 2 mm. b) hydranth with medusae buds; scale bar 0.5 mm. c)
stenoteles of polyp; scale bar 10 Jim. d) newly released medusa; scale bar 0.5 mm. e) lateral (top) and side view (below) of
tentacle bulbs; scale bar 0.2 mm. f) nmature male medusa, form with rounded bell, same scale as d). g) nematocysts of
medusa; two stenoteles, desmoneme, same discharged, stenotele of exumbrella, same discharged; scale bar 10 pm.

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Medusa buds arise from short stems in upper axil
of the tentacles. Medusa budding zone may cover
proximal two-thirds of hydrant there are occasionally
2 buds per tentacle, each with its own stem. Not only
hydranths of erect part of one colony bear medusa
buds, but also stolonal polyps from the same colony.
Nematocysts and additional observations:

a) larger stenoteles, (20-24) x (13.5-15) jim.

b) smaller stenoteles, (11-13) x (6.5-7) jim.

Caulus diameter 160-260 |im.

Colours: epidermis and tentacles transparent; gastro-
dermis variable, orange to light green, perisarc amber.

Young medusa: Medusa after liberation bell-shaped,
reaching 1 mm height and 0.8 mm diameter, jelly thin,
velum approximately half of the radius; with 8
adradial nematocyst tracks, lacking exumbrellar fur¬
rows. Manubrium not longer than half the subum-
brellar cavity and lacks nematocysts. Without apical
canal. Four radial canals ending in tentacle bulbs with
abaxial red-brown ocelli; 4 tentacles bear clasping
nematocyst clusters and a terminal spherical cluster.
Gonads not visible. Nematocysts:

a) smaller stenoteles in tentacles, (9.5-11) x (6.5-
8) jim.

b) larger stenoteles in tentacles, (12-14.5) x (9-9.5) |im.

Fig. 78. Sarsia eximia medusa; large cylindrical form; scale
bar 1 mm.

c) stenoteles on exumbrella, shaft with little or no
constriction near large spines, (11.5-13) x (8.5-
10) pm, s = 1.

d) desmonemes, discharged with 3 coils, (9-11) x (5-
5.5) pm.

Adult medusa: Bell-shaped to cylindrical, normally
3-4 mm high (2.2-7 mm range), jelly thicker at apex,
without nematocysts on exumbrella. Manubrium
cylindrical and spanning half to all of subumbrellar
height. Gonads encircle the manubrium, extending
from near its base almost to the mouth. Four marginal
bulbs with red-brown ocelli and 4 very extendible
tentacles. The abaxial side of the tentacle bulb
gastrodermis is convex, the radial canals enter
adaxially. Tentacles bear 50-100 nematocyst clasps in
indistinct spirals and one terminal cluster. Nemato¬
cysts and additional observations:

a) larger stenoteles, (14.5-17.5) x (10.5-12) pm.

b) smaller stenoteles, (10.5-13) x (7-9) pm.

c) desmonemes, (11.5-12) x (5-6.5) pm.

Egg diameter (spawned) 192 pm (n = 24, 5 animals
from 3 different dates); diameter of terminal capitae
80 pm.

Colours: manubrium light green, tentacles light
orange.

Type Locality: British Isles.

Life-cycle Observations: The hydroid is quite easy to
cultivate. In culture its tentacles may become much
longer. Medusae released from three different colonies
were raised to sexual maturity (spawning) at about
2.6 mm bell height within 18-22 days (room tempera¬
ture). Medusae thus obtained were not distinguishable
from those taken from the plankton.

Remarks on Variation: The perisarc at the base of
hydranths is normally only visible under a compound
microscope. In some colonies, however, it forms an
obvious basal cup reaching up to the most proximal
tentacles. Although these colonies also have normal
hydranths they can be mistaken for another species.
In culture, such colonies loose their cups and the
medusae grow into normal S. eximia . The occasional
cups may be a general feature of S. eximia , as Brinck-
mann-Voss (1989: fig. 5) also depicted polyps with an
indication of basal perisarc cups.

The medusa shows an even greater degree of vari¬
ation. Frequently, medusae showed a highly cylin¬
drical bell with a thickened apical jelly and flat top
(Fig. 78). The subumbrella, however, always had
curved walls. This form was first taken for another
species, namely Sarsia gracilis Browne, 1902, but the
typical juvenile forms of S. gracilis described by

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Browne and Kramp (1939) were never found, despite
the majority of all collected Sarsia medusae being
juvenile. Several other observations showed that the
cylindrical medusa is a form of S. eximia: there is a
continuum of morphological forms in between the
shapes given in figures 77f and 78; there were no
significant differences in nematocyst or egg size; both
forms interbreed easily (see Appendix). Although it
was not possible to quantify fertilisation success, most
spawned eggs developed. Crosses between sibling
species of Sarsia are possible, but viable larvae are few
(Miller 1982). Hartlaub (1907, fig. 2a) and Brinckmann-
Voss (1989) also depicted S, eximia with thickened
apical jelly. The size of the bell was normally in the
range of 3-4 mm, but exceptionally there were some
specimens reaching 6 mm, and one even 7 mm.
According to Russell (1953), S. exiwia can rarely reach
10 mm in height and the length of the manubrium,
which can be as long as the subumbrella, is variable
in this species (cf. Appendix).

Barnett (1985) recorded S. exiwia and S. gracilis
from Leigh. She mentioned that the two species were
difficult to identify because there were also inter¬
mediate forms. Having examined part of her material,
her S. gracilis is here referred to S. exiwia , being within
the range of the animals observed in Wellington.

General Remarks: Brinckmann-Voss (1989) recently
reviewed the data for species of the S. eximia group,
and further information can be found in Watson (1978)
and Kubota and Takashima (1992).

Conjne tenella was described as a polyp occurring
in Wellington Harbour on holdfasts of Macrocystis
(Farquhar 1895). His description and figures also fit
S. eximia. A problem arises from his remark that the
gonophores are sessile, but he may not have used the
term in the modern sense, meaning that no medusa is
liberated, as he did for Tubiclava rubra (see Turritopsis
nutricula). Later, Bale (1924) kept C. tenella separate
from S. eximia as Syncoryne tenella but did not clearly
state why. He thought that the medusa may not
become free, which is quite improbable as he observed
ocelli. Ralph kept S, tenella distinct from S. eximia by
stating that the former has gonophores without stems.
This contradicts Bale's description (1924) and also her
figure to some extent.

Examination of the holotype of Coryne tenella
showed that it cannot be separated from the S. eximia
material collected for this study. For correct identi¬
fication of mostcorynid polyps, the released medusae
must be raised to maturity because many Sarsia and
Coryne species cannot be distinguished in the polyp
phase alone. Farquhar's species may, therefore, never
be identified with certainty; however, its accordance
with S. eximia polyps, which are abundant in Welling¬

ton Harbour (including on Macrocystis) make it very
probable that Farquhar's sample was S. eximia . Sarsia
eximia hydroids often form very dense populations in
Wellington Harbour, particularly on floating objects
like Macrocystis , buoys or rafts. On a raft used for
mussel culture they formed colonies occupying square
metres. Sarsia eximia can also grow on ship hulls
(Millard 1975) which may explain its wide distribution.

The present samples agreed in most respects with
published descriptions of S. eximia. Only a few
characters did not fit the data of other authors. The
tentacle bulbs of the medusa were not found to be
laterally compressed as assumed by Petersen (1990),
and other studies do not show such a morphology
(e.g., Brinckmann-Voss 1989). Additionally, it was
noted that the shape of the gastrodermal chambers
depends on how recently food has been consumed.
Contrary to Petersen (1990), the tentacles of the polyp
are mostly not arranged in whorls. An irregular
arrangement of the tentacles is also given by Brine k«
mann-Voss (1989) and Russell (1953). The most signifi¬
cant difference was the egg size of 192 jum. This is in
contrast to data of Bodo and Bouillon (1968, 130 pm)
and Brinckmann-Voss (1989,113 pm). This difference
is larger than the variation observed within popu¬
lations (cf. Rees 1957, table 2; Miller 1982, fig. 2), but
unfortunately nothing is known about variations
between populations.

A similar species which could also be present in
New Zealand is the Australian Sarsia radiata von
Lendenfeld, 1884, but polyps are distinguishable from
S. eximia by their microbasic euryteles (Watson 1978).

Records from New Zealand: The hydroid is known
from Wellington Harbour (Farquhar 1895, as Coryne
tenella , this study); and Oamaru (Bale 1924). The
medusa is known from Cook Strait (Kaberry 1937);
Oamaru Harbour (Ralph 1953); marine reserve Leigh
(Barnett 1985, as S. exiwia and S. gracilis); Wellington
Harbour (this study).

Other Records: North Atlantic from America to
Europe and from Iceland to France; Mediterranean;
west coast of North America. Medusa also known
from Brazil, Valparaiso, and the N W Pacific (Millard
1975).

Sarsia japonica (Nagao, 1962) (Fig. 79a-g)

Stauridiosarsia japonica Nagao, 1962: 176, figs 1-5.

[not Sarsia japonica Maas, 1909 ~ Euphysa japonica]

Sarsia japonica: Arai & Brinckmann-Voss 1980: 21, fig. 10;

Brinckmann-Voss 1989: tables 2 & 3; Kubota & Taka¬
shima 1992: 371, figs 1-14.

Coryne japonica: Petersen 1990: 211.

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Fig. 79. Sarsia japoiiica, all afrom life, a) cultivated polyp stage with medusa buds; scale bar 0.5 mm. b) nematocysts of
polyp stage: stenoteles; scale bar 10 mm. c) newly released medusa; scale bar. d) side and frontal view of tentacle bulbs, scale
bar 0.2 mm. e) adult medusa from culture, same scale as c). f) part of tentacle with tuft of long, stiff cilia, CL; scale bar 50 pm.
g) nematocysts of young medusa: stenotele, desmoneme, exumbrellar mastigophore, same discharged; scale bar 10 pm.

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Material Examined:

1 polyp colony on holdfast of drift Macrocystis, collected
4.7.1994 at Lyall Bay Beach,Wellington.

1 polyp colony growing on oyster, collected 28.6.1994
underneath the pier restaurant, Greta Point, Evans Bay,
Wellington, 0.5 m.

1 colony growing on red sponge, collected 5.9.1994 near
NIWA, Greta Point, 0.5 m.

All above colonies were infertile but were cultivable in
plastic dishes until medusae were produced. Approxi¬
mately 30 medusae were released and most of them reared
to maturity. It takes two weeks to reach full maturity, but
gonads are easily visible after one week. Some material
deposited.

Description:

Polyp stage: Capitate hydroid colonies, mostly
stolonal, only occasionally branched, arising from
attached, ramifying stolons. Cauli longer than
hydranths (in culture). Perisarc covers stolons and
cauli, not annulated. Hydranths claviform, up to
1.5 mm. With a distal whorl of 4- 5 capitate tentacles;
below them 3-4 whorls, each with 4 capitate tentacles,
all of similar length, only slightly tapering, up to 20
gastrodermis cells in the most distal tentacles. Below
capitate tentacles a whorl of 4-5 filiform tentacles.
Medusae buds arise either singly or in groups of up
to 3 among the lowest tentacles, some of them may
also arise in the upper axil of these tentacles. Medusae
buds on relatively long, unbranched stalks. Ocelli of
medusae buds point in axial direction. Nematocysts:

a) smaller stenoteles, (12-14.5) x (7-8.5) pm.

b) larger stenoteles, (19-19.5) x (12-15) pm.

Newly released medusa: 1.4 mm high, bell-shaped,
with 8 adradial rows of nematocysts; the rows may
be rather indistinct in some specimens but the
perradial and interradial positions are always free of
nematocysts. Manubrium length one-third of bell
cavity, a slight rounded apical chamber present or
absent, gonads not developed. Four tentacles with 10-
15 nematocysts clusters and a terminal cluster of
similar size, most nematocyst clusters encircling
tentacle (moniliform condition). On the first cluster
beyond the terminal one, and additionally on 1-2 more
proximal ones, are adaxial tufts of stiff, long cilia
(visible at higher magnifications only). These tufts
move actively. Ocelli dark red. Nematocysts:

a) stenoteles in tentacles, (13.5-16) x (8.5-11) pm.

b) stenoteles from exumbrella, shaft tapering, con¬
striction at level of large spines almost absent, ca.
15 x 11 pm, s = 1.1.

c) desmonemes, (10-11.5) x (4-5.5) pm.

Adult medusa (after two weeks in culture): Up to
2.7 mm high, higher than wide, bell-shaped with apex
slightly flattened, jelly slightly thicker at apex. Dilated

velum spanning half of radius. Manubrium half to
two-thirds the subumbrellar depth, manubrium apex
slightly rounded, gonads surrounding manubrium
completely leaving only parts near mouth and apex
free. Large tentacle bulbs withabaxial ocelli and long
tentacles. Ocelli dark red. Tentacles expanded 2-3
times bell length, with 60-70 nematocyst clusters,
these, except most proximal ones, apparently encircle
relaxed tentacle (moniliform tentacle). One terminal
cluster of similar size as other the clusters. Tentacles
are mostly fully expanded and actively moving.
Several of the nematocyst clusters have an adaxial tuft
of 10-20 long stiff cilia that move actively. No exum-
brellar nematocysts present.

Type Locality: Akkeshi, Hokkaido, Japan.

Remarks: The present sample of S. japonica agrees in
most respects with the descriptions given by Nagao
(1962) and Kubota and Takashima (1992). The most
obvious difference is the presence of tufts of stiff cilia
on the tentacles. These tufts are not easily seen and
require observation at high magnifications with
suitable illumination. It may well be that they have
been overlooked by previous workers. Other differ¬
ences are the smaller size of the larger stenoteles of
the polyp and the presence of some medusae buds in
the upper axil of the tentacles. As amended by Kubota
and Takashima (1992) the stems of the medusae buds
are not branched blastostyles as given in the original
description of Nagao (1962). The length of the manu¬
brium in the present sample conforms more with the
description of Nagao (1962) than with that of Kubota
and Takashima (1992).

The nematocysts on the exumbrella of the newly
released medusae have been described as microbasic
mastigophores (Kubota & Takashima 1992), but their
figures 11-12 do not correspond to this type. The
exumbrellar nematocysts found in the present sample
were interpreted as somewhat aberrant stenoteles, an
interpretation that can also be applied to the figures
of Kubota and Takashima (1992). The shafts of these
stenoteles taper only slightly and mostly have no
constriction at the level of the large spines (Fig. 79g).
A stenotele with a similar shaft, although less deviant
from the normal type is also found on the exumbrella
of S. eximia (Fig. 77g).

Petersen (1990) used the position of the medusae
buds as one character to redefine the genera within
the Corynidae. But in S. japonica the occurrence of
medusae buds in the axils of the lowest tentacles as
well as independent of them makes the revised
classification of Petersen (1990) less applicable. The
same has been noted for the morphology of the
tentacular bulbs (Kubota & Takashima 1992). The

130

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shape of the gastrodermal chamber, however, seems
not to be a constant feature and varies with the amount
of food present in the gastric system (unpubl. observ.
for several corynid medusae).

The adult medusa of S. japonica is very similar to
S. exiwin (cf. Figs 77f, 79e). In preserved material they
may be not distinguishable, as not all specimens of S.
japonica have an apical chamber. The presence of tufts
of cilia on the tentacle is, however, a character that
might allow a clear distinction between these species.
Additionally, the tentacles of S. japonica show a
different behaviour as they are more often kept
expanded and move more actively. Unfortunately,
medusae of S. japonica are not yet known from New
Zealand waters, and it is not known whether natural
populations always have these cilia and tentacular
behaviour.

The adult medusa may reach 5-6 mm in height
(Arai & Brinckmann-Voss 1980; Kubota & Takashima
1992).

Infertile polyps of S. japonica are very similar to
Dipurena ophiogaster (cf. Figs 79a and 76d). Only the
site of medusae budding is different In D. ophiogaster
the buds are mostly located below the capitate
tentacles.

Records from New Zealand: Lyall Bay and Evans Bay,
Wellington (this study).

Other Records: British Columbia, Canada; California
(?); Japan (Kubota & Takashima 1992).

Family CLADONEMATIDAE Gegenbaur, 1857

Capitate hydroids forming stolonal or sparingly
branched colonies. Stolons ramifying, these and stems
covered by perisarc reaching to base of hydranths.
Hydranth spindle-shaped, with an oral whorl of four
to five capitate tentacles with chordoid gastrodermis,
with or without an aboral whorl of four to six slender,
solid filiform tentacles with few or no nematocysts
but with sensory hairs. Mouth with oral mucus glands
enclosed by ectodermal lips forming preoral chamber.
Medusa buds not enclosed in perisarcal film, carried
singly or in clusters distal to aboral whorl or at base
of hydranth.

Medusae with variable number of bifurcating or
simple radial canals. Stomach cylindrical, with or
without stomach pouches, mouth circular with or
without short protuberances armed with nematocysts.
Gonads on stomach wall, on subumbrellar surface or
in special brood pouch. Branching marginal tentacles
with adhesive organs, corresponding in number to,
or exceeding number of radial canals that reach ring

canal. Gastrodermis of tentacles parenchymatic.
Abaxial ocelli present (modified after Petersen 1990).

Remarks: Characteristics of genera known from New
Zealand:

Cladonema : medusa with bell high, tentacles branching
more than once.

Staurocladia : flat medusae, tentacles branching once
only.

Cladonema Dujardin, 1843

Hydroid colonial, mostly stolonal and only occasion¬
ally branching. Hydranth with a whorl of four to five
oral capitate tentacles; with or without four to five
filiform aboral tentacles. Medusa bud borne singly
on body of hydranth immediately above aboral ten¬
tacles or in similar position when aboral tentacles are
absent.

Medusa adapted for swimming and creeping. With
bifurcating and simple radial canals. Cylindrical
stomach with perradial stomach pouches and four to
six oral nematocyst clusters. Gonads surround
stomach. Branching marginal tentacles correspond in
number to radial canals that reach ring canal, each
with one to six branches ending in adhesive organ and
one to ten branches with clusters of nematocysts (after
Petersen 1990).

Type Species: Cladonema radiatum Dujardin, 1843.

Remarks: The taxonomy of the genus Cladonema is, at
present, tentative. The species limits are far from clear
and much more information is needed to delimit them.
A good review is given by Rees (1979a) — there seem
to be at least three species, namely C. radiatum
Dujardin, 1843, C. californicwn Hyman, 1947 and C.
pacificum Naumov, 1955.

In New Zealand there is only one known finding
of a Cladonema.

Cladonema radiatum Dujardin, 1843 (Fig. 80a-d)

Cladonema radiatum Dujardin, 1843: 1134; Mayer 1910: 90,
figs 53-55; Weill 1937:443, fig. 1 (cum s\jn.); Russell 1953:
105, figs 49-51; Brinckmann & Petersen 1960: 388, fig. 2;
Kramp 1959: 96, fig. 55; Kramp 1961: 57 (cum. si//;.);
Kramp 1968: 22, fig. 52; Brinckmann-Voss 1970: 76, figs
88-89, pi. 5 figs 1-2; Rees 1979a: 300; Calder 1988: 67, fig.
50 (cum syn), Hirohito 1988: fig. 12a-b.

Cladonema perkinsi Mayer, 1904: Mayer 1910: 101, pi. 9 fig.
1 .

Cladonema mayeri Perkins, 1906: 118; Mayer 1910:101, pi. 9
figs 2-3.

Cladonema novae-zelandiae Ralph, 1953: 72, fig. 20 (new
synonym).

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Material Examined:

Microslide with holotype otCladonema novaezelandiae Ralph,
1953, MoNZ Co208, collected 1949 in rockpool in Island
Bay near Fisherman's Creek, Wellington.

Microslide with paratype of Cladoneina novaezelandiae,
collected 1949 by P. Ralph, MoNZ Co209.

Microslide made by P. Ralph in 1949, held by MoNZ,
labelled: M2 Cladoneina radiatum californica (radiatum later
crossed out).

Cladoneina radiatum polyps and medusae from the Mediter¬
ranean (Banyuls-sur-Mer), life cycle observed from polyp
to medusa.

Description:

Medusa stage ( Mediterranean material): Medusa up
to 3 mm high with bell-shaped umbrella, slightly
higher than wide, jelly moderately thin, velum rather
broad. Medusa able to swim freely but mostly remain¬
ing attached. Manubrium spindle-shaped with 4-5
perradial pouch-like protuberances in its middle
region. Mouth with 4-5 nematocyst clusters. Manu¬
brium does not extend beyond umbrella margin. The
gonads completely surround the manubrium in the
region of the pouches. Radial canals may bifurcate and
up to 10 canals may reach the circular canal; the
branching pattern varies between individuals.
Number of marginal tentacles corresponds to number
of radial canals. Marginal tentacles branched with
elongated thickened bases, from the distal underside
of which grow 1-4 (up to 6 according to Weill 1937)

short tentacles with adhesive organs. The branched
upper portion of the tentacles may have up to 5 ends
with numerous nematocyst clusters alternating in
position (aboral and oral) and one larger terminal
cluster. Each tentacle base has an abaxial ocellus.
Nematocysts:

a) larger stenoteles, (13-16) x (9-10) jam.

b) smaller stenoteles, (9.5-11) x (5-8.5) |im.

c) desmonemes, (9-12) x (3.5-5) jam.

Polyp stage (Mediterranean material): Hydroid
colonies arising from attached, ramifying stolons,
occasionally with branching stems, reaching a height
of up to 2.5 mm. Perisarc smooth, terminating shortly
below filiform tentacles. Hydranths ca. 1 mm high,
with a whorl of 4-5 capitate tentacles with 7-9
chordoid gastrodermal cells. Below capitate tentacles
a whorl of 4-5 slender, filiform tentacles with a slight
terminal swelling. Hypostome with an ectodermal
preoral chamber. Medusae buds naked, distal to fili¬
form tentacles. Nematocysts:

a) stenoteles, (11-17) x (8-10) |im.

b) mastigophores, rare or absent, in stolons only, (10-
12) x (3.5-4) pm.

Type Locality: First described from the Mediterranean
Sea.

Remarks: The nominal species Cladoneina novae -

Fig. 80. Cladoneina radiatum , from live Mediterranean material, a) young medusa; scale bar 0.5 mm. b) nematocysts of
medusa: two types of stenoteles, desmoneme; scale bar 10 pm. c) polyp stage; scale bar 0.2 mm. d) nematocysts of polyp:
stenotele, rare mastigophore; same scale as b).

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zelnndine was erected by Ralph (1953) based on
medusae found near Wellington. Her descriptions are
limited and the reasons for creating a new species are
not apparent The original type material of C. novae-
zelandiae was examined for this study. The samples
comprise a holotype and a paratype specimen
mounted onto microslides. The morphology of both
animals is only very poorly preserved, but permits
observation of some characters valuable for a more
precise taxonomic evaluation. The most important are
as follows: manubrium shorter than bell height,
gonads cover middle part of manubrium, 9 tentacle
bulbs with much-branched tentacles with up to 6
adhesive branches and up to 5 branches with nemato-
cyst clusters. Ralph (1953) related her species to C.
californicum, which, however, is not the case. Cladonema
californicum has a manubrium that protrudes through
the velar opening, has no branching radial canals, and
far fewer tentacle branches (Rees 1979a). As the gonads
in Ralph's medusa do not cover the whole manubrium,
C. pacific tun (syn. C. uchidai) can also be ruled out (cf.
Hirohito 1988). Ralph's description of C. novnezelnndiae
(1953), her figures, and the results of the re-examin¬
ation of the types match rather well the description of
C. mayeri given by Mayer (1910, p. 101), especially the
9 tentacles, each with up to 6 adhesive branches per
tentacle. Weill (1937) examined the variability of
Cladonema at Bermuda and concluded that C. mayeri
is a variant (and junior synonym) of C. radiation only.
Mayer (1910) also considered this as a possibility. For
these reasons, C. novaezelandiae is here synonymised
withC. radiation . However, further information on the
New Zealand population, especially its polyp stage,
is needed to confirm this. In order to facilitate future
recognition of differences or similarities of the local
population to the population of C. radiation from the
type locality, the descriptions and figures give above
were made from Mediterranean material.

Records from New Zealand: Known only from
Wellington (Ralph 1953).

Other Records: Western Atlantic; Europe; Medi¬
terranean; Japan (Calder 1988; Hirohito 1988).

Staurocladia Hartlaub, 1917

Hydroids, where known, stolonal or on erect,
nonbranching cauli. Hydranth with an oral whorl of
capitate tentacles, with or without four to five aboral
filiform tentacles. Medusa buds borne singly on body
of hydranth immediately above aboral tentacles or in
similar position when aboral tentacles are absent.
Medusa adapted for walking. Umbrella with continu¬

ous, broken, or no nematocyst ring along margin. With
six to eleven radial canals, some shortly bifurcating
distal to manubrium. Manubrium with circular mouth
with or without five to six knobs armed with nemato-
cysts. Gonads surrounding stomach or developed on
protrusions from stomach. With up to 60 bifurcating
marginal tentacles, one upper branch with several
nematocyst clusters, one lower branch with adhesive
organ. Asexual reproduction through fission, or
budding of medusae from umbrella margin.

Type Species: Staurocladia vallentini (Browne, 1902).

Remarks: The above diagnosis was modified after
Petersen (1990) to accommodate Staurocladia welling -
toni n.sp., which has sessile, stolonal polyps without
cauli and also no nematocyst ring in the medusa. The
latter character reduces the distinguishing characters
of Staurocladia and Cladonema. The assignment of these
two genera to two separate families Cladonematidae
and Eleutheridae thus becomes even less reasonable
than before (see Petersen 1990). Another character is
the structure of the gonads which seem not to be
located on manubrial protuberances in all species. In
the original description of Staurocladia hodgsoni
(Browne, 1910), the gonads were described as com¬
pletely surrounding the manubrium in a continuous
ring, with seven or eight swellings each containing a
single large egg. All species of Staurocladia have been
reviewed in tabular form by Bouillon (1978a).
Characteristics of the New Zealand species:

S. vallentini: tentacular nematocyst clusters of medusa
on oral and aboral sides, polyp with filiform
tentacles.

S. wellingtoni : tentacular nematocyst clusters of
medusa lateral, polyp without filiform tentacles
and with desmonemes.

Staurocladia vallentini (Browne, 1902) (Fig. 81a-e)

Eleutheria vallentini Browne, 1902: 279.

Cnidonetna capensis Gilchrist, 1918: 509, pi. 30.

Staurocladia vallentini: Browne & Kramp 1939: 274, pi. 14,
figs 3-4, pi. 15 fig. 4, pi. 19, fig. 2; Kramp 1959: 98, figs 58-
59; Kramp 1961: 62; Millard 1975: 57, fig. 23D-G; O'Sulli¬
van 1982: 29, fig. 11, map 10; Hirohito 1988: 42, figs 12c-e.

Cnidonewn vallentini: in part Ralph 1947: 414, pi. 35, figs 1,
2,3, 4A, not others.

Cnidonema hodgsoni: in part Ralph 1953: fig. 16, polyp.

[not Staurocladia hodgsoni (Browne, 1910)].

Material Examined:

Ca. 30 medusae collected from Cystophora , other macroalgae,
and bryozoans, from January to October 1994 at Te Rae-
Raekaihau Point, Houghton Bay, and Island Bay, Welling-

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ton, all sizes from 6-24 tentacles, none fully mature,
intertidal to 13 m. Many were also cultivated but they are
rather difficult to feed.

Microslide prepared by P. Ralph, held at MoNZ. Label 1
(older): Cnidonemn vallentini , 1946, Ml. Label 2: Staiiroclndia
hodgsoni Browne, Ml. The slide contains two medusae in
poor condition. The larger one is 1.4 mm in diameter with
31 tentacles with 1-2 accessory nematocyst clasps in aboral
position, none in oral position, an aboral ring of nemato-
cytes tissue, and 4 or more gastric pockets (gonads?) fill

the subumbrella. The smaller one is 0.5 mm in diameter,
with 7 tentacles but originally more, only a terminal
nematocyst cluster on the tentacles and there may be an
aboral nematocyst ring may be present.

Description:

Medusa stage: Crawling medusa unable to swim
freely, bell much reduced, diameter up to 1.5 mm and
only half as high. Manubrium pear-shaped, almost
filling subumbrella; oral part of manubrium tubular

Fig. 81. Staurocladia vallentini , from life except e). a) medusa, not fully mature; scale bar 0,5 mm. b) cnidome of medusa:
three sizes of stenoteles, desmoneme; scale bar 10 pm. c) lateral view of tentacle showing the aboral and oral position of the
nematocyst clusters (shaded), CL; scale bar 0.2 mm. d) oral view of medusa, m manubrium, mb medusa bud, nr nematocyst
ring, rc radial canal, te tentacle, v velum; CL, same scale as c). e) polyp stage with medusa bud, redrawn from Ralph (1953)
with permission of Victoria University Press, same scale as a).

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and very extensible, without nematocyst clusters
around mouth. Gonads bulge from manubrium walls.
Velum very broad, reaching manubrium. Between
velum and bell margin a continuous ring of thickened
epidermis containing many nematocysts. This region
buds off new medusae in immature animals. Mostly
1 (6-11) short, unbranched radial canals issue from
manubrium. Circular canal very broad. Up to 30
branched tentacles, each with an abaxial ocellus at its
origin. Lower branch of tentacles with a suckerpad
used to adhere to substratum, the upper and longer
branch with a terminal knob of nematocysts and 2- 3
aboral and 0-1 aboral nematocyst cluster.
Nematocysts:

a) larger stenoteles, (19-25) x (12-16) pm.

b) medium sized stenoteles, (14-15.5) x (8.5-10.5) pm.

c) smaller stenoteles, (11-13) x (7-9) pm.
e) desmonemes, (10-11) x (4.5-5.5) pm.

Colour: pink to orange, very opaque.

Polyp stage (after Gilchrist 1918 and Millard 1975):
Hydroids colonial, hydranths born on slender hydro-
cauli arising directly from a tubular hydrorhiza,
hydranth length up to 1.5 mm, slender at base and
increasing in diameter distally, with 3-4 capitate oral
tentacles and 4-6 filiform aboral tentacles. Hydro-
caulus reaching 2 mm in length, covered with thin
perisarc. Medusae buds develop slightly above level
of filiform tentacles, with about 6 bifurcating tentacles
at liberation, the aboral branch with a terminal
nematocyst cluster.

Type Locality: Falkland Islands.

Remarks: Staurocladia valient ini has been known from
New Zealand since Ralph's 1947 paper, in which she
described various stages of medusae and polyps found
in her aquarium and the sea. It is not apparent that
Ralph followed the development of single medusae
— which is rather difficult — but most probably she
assembled various sizes of medusae into a develop¬
mental pattern. She described some unusual develop¬
ments such as the shifting of the position of the
tentacular nematocyst clusters and also variations in
the gonads. Such features are atypical and were not
observed in the present study. From re-examination
of part of her material and much new living material
it became evident that she had two different species
before her, S. valient ini and S. wellingtoni, a new species
described below. A polyp depicted in Ralph's work
of 1953 as Staurocladia hodgsoni (plate III, fig. 16A) is
most probably the polyp of S. vallentini. Its medusa
bud has six tentacles and only terminal nematocyst
clusters. The figure and her description of the polyp
(Ralph 1947) differ from other descriptions of S. vallen-

tin i only in the more distal position of the medusa bud
(cf. Gilchrist 1918, Hirohito 1988). Unfortunately no
polyps could be found in this study. The medusae of
the available material agreed well with other
descriptions. The only observed difference was the
higher number of radial canals, which is rather vari¬
able in other populations too. The youngest observed
medusae had six radial canals.

Staurocladia wellingtoni differs radically from S.
vallentini at all stages. The main differences are: size,
colour (orange versus yellow-brown), position of
tentacular nematocyst clasps, presence of a nematocyst
ring, absence of oral nematocystclusters, and presence
of filiform tentacles in polyp. It was interesting to note,
however, that both were often found together on the
same algal frond.

Records from New Zealand: Wellington area (this
study), Leigh (Barnett 1985).

Other Records: Falkland Islands; Australia; South
Africa; Japan (Hirohito 1988); Bermuda (Kramp 1959).

Staurocladia wellingtoni n. sp.

(Figs. 82a-g, 83a-d, cover)

Staurocladia vallentini: in part Ralph 1947: 417, fig. 4B.
Staurocladia hodgsoni : in part Ralph 1953: 70.

Material Examined:

Four slides prepared by P. Ralph, held at MoNZ Welling¬
ton, labelled 11 Staurocladia hodgsoni July 1952 from Aq.
Tank 11 ; 8 medusae total.

Ca. 60 living adult and many juvenile stages from 16-tentacle
stage on, collected 30.1.94, 11.3.94, 26.4.94, and 25.8.94,
on macroalgae Cystophora torulosa and C. retroflexa at 0.5-
2 m, western side of Te Raekaihau, Wellington. One male
medusa from first batch was selected as holotype H-650.
Part of the remaining material is deposited as paratype P-
1080.

A colony with 5 polyps (2 with medusae buds) growing on
stone from western side of Te Raekaihau Point, collected
11.4.94, at 1-2 m, cultivated in aquarium until 23.5.94 when
polyps were discovered. One of the two medusae buds
was released and the medusa cultivated to the 24-tentacle
stage. Polyps and especially medusae were easy to
cultivate. The medusa was kept in a small petri dish in
5 ml sea water and fed daily with several Artemia nauplii.
Medusa deposited.

Description:

Adult medusa: Crawls on macroalgae and unable to
swim freely. Umbrella a shallow bell, much wider than
high, diameter up to 4 mm without tentacles, the jelly
thin but noticeable. Velum broad, fitting closely
around the manubrium which may protrude from the

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Fig. 82. Staurocladia wellingtoni n.sp., all fromlife. a) adult medusa from western side of Te Raekaihau, Wellington; scale
bar 1 mm. b) aboral view oLbell of female medusa, no tentacles shown, same scale as a), c) oral view of manubrium and
gonad sacs of a male medusa, note the six nematocyst knobs (me) on lip, same scale as a), d) aboral view of bell of a young
medusa from nature, 16-tentacle stage, same scale as a), e) side view of manubrium of medusa depicted in d), shaded area is
pigmented black, note presence of nematocyst knob on lip margin, same scale as a), f) side view of manubrium of an adult
medusa, the mouth knobs (me) continue as ridges along the manubrium; scale bar 0.2 mm. g) nematocyst of adult medusa,
from right: four sizes of stenoteles, desmoneme and discharged desmoneme; scale bar 10 pm. Abbreviations: go gonads, mn
manubrium, me mouth knobs, ms manubrial sacs, rc radial canals.

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subumbrella. The upper part of the tubular manu¬
brium with 6, rarely 5, large mobile protrusions (sacs)
covered by the gonads. Female gonads contain a large
number of small eggs (diameter 74 pm, spawned).
Mouth rim is lined by 6 conspicuous knobs of nemato-
cyst clusters that continue over the outside of the
manubrium as gentle ridges without nematocysts.

Long radial canals originate at apex of manubrium;
normally 8 complete radial canals, 2 pairs of which
may fuse near the manubrium. This branching pattern
variable. Between the complete radial canals, there
are 2-4 centripetal, incomplete radial canals origi¬
nating from the circular canal Radial canals are darkly
pigmented. The pigmentation of the complete canals

Fig. 83. Staurocladia wellingtoni n.sp., all from life, a) CL drawing of tentacle of adult medusa viewed from aboral,
pigmented region in gastrodermis is shaded; scale bar 0.2 mm. b) CL drawing of tentacle bulb region of an adult medusa
from oral, the arrow indicates a nematocyst, no continuous ring of nematocysts is present; scale bar 0.2 mm. c) aboral view
of a one day old medusa released form polyp shown in d); scale bar 0.2 mm. d) polyp stage with medusa one-day before
liberation, same scale as c). e) nematocysts of polyp stage, from right: three sizes of stenoteles, desmoneme and discharged
desmoneme; scale bar 10 pm. Abbreviations: cc circular canal, rc radial canal, te tentacle.

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fades towards the centre and then becomes a white
star- or hexagonal-shaped disk on top of stomach base.
The complete radial canals are not in phase with the
gonads. Up to 55 marginal tentacles, each up to 3 mm
long with a red abaxial ocellus. The tentacles are
bifurcated into a lower branch ending in an adhesive
organ used for walking and adhering, and an upper
branch with nematocyst clusters. The upper branch
longer than the lower one, bearing a terminal spherical
cluster and up to 6 lateral clasps of nematocyst
alternating on each side. Tentacles hollow, the lumen
lined by dark pigment. No nematocyst ring along the
bell margin. Nematocysts:

a) stenoteles size class 1, (28-30.5) x (20-21) jim.

b) stenoteles size class 2, (17.5-24) x (13-15) |im.

c) stenoteles size class 3, (9.5-14.5) x (7-9.5) Jim.

d) stenoteles size class 4, (7-9) x (5.5-6.5) |im.

e) desmonemes, discharged with 3-4 coils with
bristles, (10.5-14.5) x (4-6) |im.

Colours in life: gonads of female brown-yellow
opaque, in male olive-green transparent, both gonads
covered with black pigment towards the centre; lumen
of tentacles and radial canals dark brown to black,
apex of manubrium white; sucker pads of lower
tentacle branch light yellow.

Newly released medusa: Bell diameter 0.5 mm,
umbrella flat, with 8 branching tentacles, lower branch
with adhesive organs, upper branch with a terminal
spherical nematocyst cluster and a pair of lateral
clasps. Eight ocelli. No nematocyst ring, no manubrial
sacs, with 6 small nematocyst clusters on mouth
margin. Eight short radial canals, medusae buds
lacking (also during later development).

Colour: pink.

Growth table (time in days):

1: Newly released medusa (see description and Fig.
83c).

3: New tentacle buds grow.

5: Bell grows, radial canals lengthen.

6: Stomach turns white, four pairs of lateral clasps.

8: 16 tentacles, now the same morphology as the
youngest stages observed from nature (cf. Fig.
82d).

14: Bell diameter 1 mm, 2-3 pairs of lateral clasps.

17: Four pairs of lateral clusters, 16 tentacles, stomach
sacs grow.

24: 24 tentacles, gonad tissue starts development, bell
1.8 mm diameter, centripetal canals start to grow.
53:24 tentacles, bell 3 mm diameter, gonads well-
formed.

Polyp stage: Hydroid colonies arising from attached,
ramifying stolons. Hydranths arise directly from
stolons and have no caulus. Hydranths spindle¬

shaped, up to 1.2 mm high, with conical hypostome.
With 4-5 oral capitate tentacles containing 8-10
chordoid gastrodermis cells. Filiform aboral tentacles
absent. Medusa bud in middle of hydranth body, not
covered by perisarc. Eight incipi-ent tentacles of
medusa visible from an early stage of development
Hydranths are very contractile and rather sensitive to
stimuli.

Colour pink-orange.

Nematocysts:

a) stenoteles, very variable in size, (4.5-17.5) x (4-
13) pm.

b) desmonemes, also in tentacles, with ropy thread in
nondischarged capsules, with bristles on inside of
coils, discharged with 4 coils, (11.5-14.5) x (4-
5.5) pm.

Type Locality: Subtidal, western side of Te Raekaihau,
Wellington, New Zealand.

Etymology: The species name refers to the type locality
near Wellington. With this naming the author also
wishes to express his deep appreciation for the city of
Wellington which made him feel really at home.

Remarks: Staurocladia wellingtoni is quite a conspicuous
medusa which can be collected only by screening
through Cystophora algae. In January 1994 it was
abundant, and there were often more than four
animals per plant. It was interesting to note that the
species occurred together with the other crawling
medusa Stciurocladici vallentini, which, in contrast to S.
wellingtoni was not mature at this time. The co¬
occurrence may be incidental, as Cystophora seems to
be the preferred substratum for epibiontic cnidarians
compared to other macroalgae in the intertidal. At that
time also the densities of the brooding anemone
Cricophorus nutrix and of thecate hydroids appeared
much higher on Cystophora than on other algae.

According to Bouillon (1978a) there are only three
species of Staurocladia with several lateral nematocyst
clusters on their tentacles. Of these, S. hodgsoni
(Browne, 1910) is the only one that has no complete
nematocyst ring and is the one most closely related to
S. wellingtoni . Due to its occurrence in Antarctica, S.
hodgsoni has not often been examined and reinvesti¬
gation with detailed comparisons to S. wellingtoni
would be valuable. Staurocladia wellingtoni differs from
S. hodgsoni (Browne, 1910) in the following details: it
is at least twice as large; has up to twice as many ten¬
tacles; completely lacks a nematocyst ring; has six
nematocyst clusters on mouth margin; long radial
canals compared to very short ones; the gonads are
on six regular, large protrusions of the manubrium;
and the eggs are considerably smaller. In S, hodgsoni,

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the gonad also forms swellings, but they are irregular,
immovable and mainly result from the large eggs. The
structure of the mouth rim in S. Iiodgsoni is not very
clear. Browne (1910: 28) described the mouth as small
and circular, but in his figure 3 of plate 3, six knobs
appear to be present. The extent of the incomplete
nematocyst ring in S. Iiodgsoni is also not clear. It may
be very similar to that inS. wellingtoni . In S. wellingtoni
there are only some nematocysts in the adaxial sides
of the tentacular bulbs, a condition also seen in many
other Anthomedusae (Fig. 83b).

Ralph (1947) noted the presence of S. wellingtoni
(as S. vallentini); most of her material (from Island Bay,
Wellington) was actually the latter species, in which
she observed the polyp and medusa stages. Ralph's
(1947) description and figures make it obvious, how¬
ever, that she also had S. wellingtoni in her material.
The only material left predating the publication date
1947 is a slide from 1946 with two S. vallentini (see
under this species). The occurrence of two species
would explain her difficulty in reconciling the different
morphologies. In her 1953 key she referred all of her
material to S. Iiodgsoni, but gave no figure of the
medusa. The illustrated and described polyp,
however, is S. vallentini.

The polyp of S. wellingtoni is unique in having
desmonemes (Fig. 83e). They are also present in the
capitae of the tentacles. Polyps of the Cladonematidae
and many other Capitata like the Corynidae do not
have desmonemes in the polyp stage. This lack has
been interpreted as a good synapomorphy for several
families by Petersen (1990). But other species within
these groups may also prove to have desmonemes see
(e.g., Coryne sp. 1).

The presence or absence of filiform aboral tentacles
is a somewhat problematic character. In many species
of Corynidae with filiform tentacles these are present
only in cultures and rarely in material from nature
(Brinckmann-Voss 1970). Only one colony of polyps
of S. wellingtoni could be examined, but they were from
a laboratory culture and lacked filiform tentacles, S.
wellingtoni polyps almost certainly always lack fili¬
form tentacles. The newly released medusa is easily
identifiable because it already has several lateral
nematocyst clusters, lacks a nematocyst ring, and has
six incipient nematocyst clusters on its mouth margin.
Other distinguishing characters of newly hatched
medusae of S. vallentini are eight tentacles as opposed
to six, and the absence of medusa buds in later
development (Figs 82d, 83c-d).

The stenotele 3 and 4 of the medusa belong to the
same, variable class. The stenoteles of the polyp are
very variable in size and are not classifiable in sub¬
classes.

Records from New Zealand: Known only from the
Wellington south coast.

Family SOLANDERIIDAE Marshall, 1873

Hydroids forming large branching colonies. With
chitinous internal anastomosing skeleton formed by
coalescence and modification of adjacent hydrocauline
tubes. Coenosarc covering entire colony and pene¬
trating skeletal interstices. Flydranths borne over
whole surface of colony, uniform in structure, cylin¬
drical; with single circlet of capitate tentacles around
mouth and numerous similar tentacles dispersed over
body. Gonophores where known cryptomedusoid or
eumedusoid sporosacs arising directly from the
coenosarc.

Type Species: Solanderia gracilis Duchassaing & Michel-
in, 1846.

Remarks: The above diagnosis is taken from Bouillon
and Cornelius (1988) and only slightly modified to
include the cryptomedusoid gonophores of Solanderia
gracilis Duchassaing & Michelin, 1846 (see Wedler &
Larson 1986).

Bouillon et ai (1992b) recently revised the family
and included two genera: Solanderia and Chitina.
Chitina was re-established by Bouillon and Cornelius
(1988) and supposedly differed from Solanderia in
having bushy colonies and hydranths confined to the
hydrocladia. As outlined below, however, this generic
separation cannot be maintained because the bushy
appearance is not a constant and characteristic feature
of S. ericopsis .

Solanderia Duchassaing & Michelin, 1846
Synonym: Chitina Carter, 1873
Diagnosis as for the family.

Type Species: Solanderia gracilis Duchassaing & Miche¬
lin, 1846.

Remarks: Only Solanderia ericopsis (Carter, 1873) is
known from New Zealand.

Solanderia ericopsis (Carter, 1873)

(Fig. 84a-g, frontispiece)

Chitina ericopsis Carter, 1873:13; Bouillon & Cornelius 1988:
1551, figs 1-5, 7.

Solanderia niisakinensis : Wineera 1968: 2, figs 1-2, pis 1-3.
[Not Solanderia niisakinensis (Inaba, 1892)]

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Material Examined:

1 colony from Goat Island, Leigh, 1 m, collected 9,8.1991,
examined alive, fertile, ca. 150 mm high, fan-shaped.

3 colonies from Barretts Reef, Wellington Harbour entrance,
11 m, collected 5.2.1994 and 13.12.1994, examined alive,
fertile, both fan-shaped but one with slightly bushy ends,
170-200 mm high.

2 small fragments fromNZ01StnC814,37°40' S,178°56.4' E,
194 m, 25.2.1962, polyps badly preserved.

1 colony from NZOI Stn 169, 36°11.2 r S, 175°17.7' E, 23 m r
14.5.1975, collected by SCUBA, size 120 x 80 mm, slightly
bushy although bent by preservation.

1 colony from NZOI Stn 1357,35°27.8' S,174°44.1' E, 0-10 m,
20.11.1977, 100 x 100 mm, hydranths well-preserved
slightly bushy due to bending in jar, even main stem bears
hydranths, fertile male.

1 colony from NZOI Stn 1378, 34°09.5' S, 172°08.7' E, 0-24 m.
collected 23.11.1977, by SCUBA, 140 mm high, no
hydranths left, fan-shaped.

Fig. 84. Solanderia ericopsis. a) colony from Barretts Reef, Wellington; scale bar 2 cm. b) terminal branch with hydranths
and gonophores, colony from Leigh; scale bar 1 mm. c) young gonophore from live material; scale bar 0.2 mm. d) mature
male gonophore with gametes partly spent, same scale as c). e) skeleton from main stem, CL, same scale as c). f) skeleton of
distal branches, longitudinal elements dominate, CL, same scale as c). g) nematocysts: stenoteles of two sizes; scale bar
10 |im.

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1 colony from NZOI Stn J673, 36*26.37' S, 175°45.60 r E, 2
m, 7.9.1974, 130 x 60 mm, dried sample, strictly fan¬
shaped.

Several fused colonies from NZOI Stn J674, 36°41.85’ S,
175°55.20’ E, 20-30 m, collected by SCUBA 8.9.1974, up to
180 mm, main stem fascicled, slightly bushy, bent into
jar, a few gonophores present even on main branches.

2 colonies from NZOI Stn J698, 37°49.8' S, 176°51.8' E, 6-
10 m, collected by SCUBA 10.7.1974, size 77 x 60 and
50 x 30 mm, soft tissues deteriorated, larger colony fan
shaped but bent, smaller one slightly bushy but also bent.

1 colony from NZOI Stn J705, 37°16' S, 176°51' E, 190 m,
collected 11.9.1974, up to 200 mm, highly bushy and
chaotic colony, attached to old stem of antipatharian or
gorgonian, most hydranths lost, with gonophores.

1 colony held by MoNZ, collected 19.9.1970, White Rock,
Wairarapa, 5 m, 120 mm high, infertile, bushy cylinder,
but bushiness mostly confined to terminal branches.

Description: Very large hydroid colonies, 5-50 cm in
height, mostly fan-shaped but part of colony or whole
may have a bushy appearance. Colony composed of
one or several fascicled stems with a diameter of up
to 25 mm, from which hydrocladia arise at irregular
intervals; yellow to light brown in colour. Branches
bifurcating several times in succession, resulting in
gradually finer and more delicate branches. When
there are several stems these may fuse; branches only
rarely anastomose. Main stems issuing from a well-
developed meshwork of root-like processes. Skeleton
internal, penetrated by and covered with coenosarc.
Skeleton of larger rootlets, main stems and principal
branches a dense network of chitinous fibres, surface
fibres thinner than deeper ones, openings round to
oval (Fig. 84f). Skeletal network of finer branches less
closely reticulate, more fenestrate, predominantly and
distinctively with longitudinal elements visible even
in living material. Longitudinal fibres joined by rather
well-spaced transverse connections. Peripheral longi¬
tudinal fibres forming gentle protrusions from surface
thus delimiting grooves of coenosarc of varied length
separated by ridges of skeleton (Fig. 84b). Hydranths
1.2-2 mm high, tubular to club-shaped with rounded
hypostome, with an oral whorl of 3-5 capitate tentacles
and further below them 12-16 scattered capitate ten¬
tacles, hydranths more numerous towards terminal
ramifications, rare on main stems. Hydranths arise
in all planes from superficial coenosarc and are not
accompanied by hydrophoral structures. If occasion¬
ally flanked by shallow triangular elevations of the
longitudinal skeletal elements these do not occur on
both sides of the hydranth. Gonophores remain fixed
as sporosacs, with 4 radial canals ending in small
bulbs, with circular canal and spadix (eumedusoids).
Size of gonophores up to 1.1 mm, oblong to spherical,
with a short stem. Nematocysts:
a) larger stenoteles, (15-18) x (12-15) pm.

b) smaller stenoteles, (6.5-9.5) x (4.5-7) pm.

Type Locality: New Zealand (exact locality unknown).

Remarks: With the exception of the generic characters,
all the examined material agrees perfectly with the
description of Chitina ericopsis given by Bouillon and
Cornelius (1988). There can be little doubt that they
belong to the same species, even more so as their
samples were from Wellington Harbour. However,
the present material shows that the generic distinction
of Chitina as having a bushy growth cannot be
maintained. The colony shape can be bushy, but more
often it is strictly fan-shaped. Both branching patterns
may even occur in the same colony. It could be that
the type of growth is determined by the prevailing
currents. In addition, it is apparent from Bouillon and
Cornelius (1988, figs 1-2) that their material is bent
and folded, probably caused by stuffing these large
colonies into smaller jars for preservation. This can
also produce a bushy appearance as was observed
during this study with fresh material. The other
character that supposedly separates Chitina from
Solanderia is the absence of hydranths from the main
branches. With more and living material examined
this character too was seen to be invalid. Hydranths
may even occur on the main trunks, although not
frequently. The only way to distinguish the two genera
therefore, would be the potential or bushy growth in
Chitina. However, this seems to be insufficient to
warrant generic distinction and Chitina is here synony-
mised with Solanderia.

From the figures given by Wineera (1968) it is quite
obvious that he also had S. ericopsis and not S. misa-
kinesis. Therefore, contrary to Bouillon et al. (1992b),
the latter species is not known from New Zealand.

The gonophores of S, ericopsis are described here
for the first time and they resemble the gonophores of
many other species in the genus. Solanderia species are
rather difficult to separate and the number of species
has been drastically reduced by Bouillon et al. (1992b).
Solanderia ericopsis , especially in its fan-shaped form,
is rather similar to S. secunda (Inaba, 1892). Although
the latter has hydrophores, they may often be reduced,
but in at least some colonies they are generally present
(Bouillon et. al. 1992b). In contrast to this, all examined
specimens of S. ericopsis lacked any hydrophoral
structures. The only other similar species of Solanderia
that completely lacks hydrophores is S. gracilis. It is,
however, distinct from S. ericopsis in having crypto-
medusoids instead of eumedusoids (see Wedler &
Larson 1986), being purple instead of yellow, and
having hydranths in one plane only.

Solanderia ericopsis forms very large, conspicuous
colonies which are often noted by divers. The colonies

141

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may even occur in rather shallow water. During long¬
term monitoring of defined rock areas around the Poor
Knights Islands, Dr C. Battershill was able to observe
one colony of S. ericopsis for more than 15 years (pers.
comm.). During this time the colony reached more
than 50 cm in height (frontispiece). Apparently, S.
ericopsis forms unusually long-lived colonies.

Records from New Zealand: Rather frequently found
around the North Island; see under material examined.

Other Records: Not known outside of New Zealand.

Family PENNARIIDAE McCrady, 1859

Colonial hydroids with tubular, ramified hydrorhiza
and upright, pinnately branched hydrocauli bearing
polyp cauli on upper side only. Perisarc tubular, thick,
and firm. Hydranths terminal, clavate to pear-shaped,
each with an aboral whorl of filiform or slightly capi¬
tate tentacles, an oral whorl of short capitate tentacles,
and one or more distinct or indistict whorls of short
capitate tentacles between.

Gonophores borne just above aboral tentacles,
either liberated as short-lived medusae or remaining
attached to hydranth as eumedusoids. Gonophore
elongated with thin mesogloea, manubrium short,
mouth lacking. Radial canals four. Tentacle bulbs four.
Tentacles absent or rudimentary. Ocelli present or
absent. Gonads surrounding manubrium. Only one
genus: Pennaria .

Pennaria Goldfuss, 1820

With the characters of the family.

Type Species: Pennaria disticha Goldfuss, 1820.

Remarks: At present, it is not clear whether Halocordyle
or Pennaria should be used to name this genus (Garcia-
Corrales & Aguirre 1985; Calder 1988; Gibbons &
Ryland 1989). Following the arguments of Gibbons and
Ryland (1989) and for the sake of stability of nomen¬
clature the name Pennaria is used in the present study.
Only one species is nown from New Zealand.

Pennaria disticha Goldfuss, 1820 (Fig. 85a-c)

Pennaria disticha Goldfuss, 1820: 89; Brinckmann-Voss 1970:
40, text-figs 43, 45-50; Gibbons & Ryland 1989: 387, fig.
5.

Halocordyle disticha : Millard 1975: 41, fig. 16C-G; Garccia

Corrales & Aguirre 1985: 85, figs 1-3 (cum syn.), Calder
1988: 57, figs 43-45 {cum syn), Oestman et al. 1991: 607,
figs 1-18; Hirohito 1988: 28, figs 9a-d, pi. 1, fig. C (cum
syn.)

Pennaria australis Bale, 1884: 45; Trebilcock 1928: 1; Ralph
1953: 70, figs 14, 21.

Material Examined:

Several colonies from on underside of raft near Devonport
wharf, Auckland, collected and preserved 13.2.1994, size
2-5 cm, one colony with medusae buds. Material
deposited.

NZOI collection Z1060, from unknown port in NZ, labelled
Pennaria australis , polyps lost.

Description: Hydroid colonies forming branching
feather-like stems up to 50 mm high, arising from
attached ramifying stolons. Growth monopodial with
terminal hydranths. Main axis often curved, not
fascicled, 0.15-0.25 mm thick, thicker in middle part,
annulated in more or less regular intervals distal to
insertion of hydrocladia (side branches). Main axis and
hydrocladia deeply black, then colour fading to clear
more distally. Hydrocladia up to 16 mm long, longest
cladia in middle of axis, originating approximately 1.5
mm apart from each other on alternate sides of main
axis. The hydrocladia of one side form an angle of
about 140° with those from the other side and are
slightly curved. They bear either hydranths on pedicels
(ramuli) or secondary hydrocladia with hydranths
again on pedicels, spacing 1.5 mm. The main axis and
the hydrocladia bear a terminal hydranth. Pedicels
(ramuli) of hydranths originate on upper side of
hydrocladia, with annulation at their base, younger
ones without annulation. Hydranths are spindle- to
pear-shaped, up to 1 mm long, with an aboral whorl
of about 12 long, filiform to slightly capitate tentacles
and up to 16 short capitate oral tentacles scattered on
distal half of hydranth. Nematocysts of aboral tentacles
mainly on aboral side and terminally (= semifiliform
according to Petersen 1990). Hypostome dome¬
shaped. Gonophores are degenerate oblong medusae
arising on short stems just above aboral tentacles, with
gonads encircling manubrium, 4 radial canals and
circular canal, 4 marginal bulbs, without ocelli or
tentacles. Nematocysts:

a) stenoteles size class 1, in capitate tentacles, (31-
36) x (17-21) iim.

b) stenoteles size class 2, (15.2-16) x (10.5-12) pm.

c) stenoteles size class 3, (8-10) x (6-7.5) pm.

d) stenoteles size class 4, (6.5-8) x (4.5-6.5) pm.

e) microbasic heteronemes with inclusion body
(mastigophore?), in aboral tentacles, not seen
discharged, (11-14) x (5-7) pm.

f) microbasic heteronemes without inclusion body
(mastigophore?), rare, only one undischarged seen,
11 x 5 pm.

142

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g) basitrichous isorhizas, (7-9) x (2.5-3) pm.

h) desmonemes, (5-5.5) x (3-4) pm.

Type Locality: Gulf of Naples, Mediterranean.

Remarks: Pennaria disticha is a large, conspicuous
hydroid which is easy to detect. The species name
australis was erected for animals with a paucity of
annulation of the ramuli. This is, however, a variable
character and all modem authors treat P. australis as
a synonym of P. disticha.

In the examined samples, no release of medusae
was observed as the colonies were kept alive for few
hours only. The sizes of the nematocysts of the present

sample agree quite well with the data of Calder (1988).
There are some differences from the data of Oestman
et al. (1991). They did not include the basitrichous
isorhiza (haploneme) seen by Calder (1988) and also
in the present samples. There is also some discrepancy
concerning the types of the heteronemes (euryteles
versus mastigophores). Unfortunately, the present
data do not allow a conclusion on that matter.

Records from New Zealand: Auckland Harbour
(Trebilcock 1928; Ralph 1953; this study).

Distribution: Circumglobal in tropical to warm-
temperate waters (Gibbons & Ryland 1989).

Fig. 85. Pennaria disticha from preserved material, a) Colony form; scale bar 5 mm. b) Part of hydrocladium with two
hydranths bearing medusae buds of different stages, at right almost mature; scale bar 0.5 mm. c) Nematocysts, clockwise
from right: four size classes of stenoteles, rare heteroneme without inclusion (mastigophore?), heteroneme with inclusion
body (mastigophore?), basitrichous isorhiza and same discharged, desmoneme; scale bar 10 |im.

143

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ACKNOWLEDGMENTS

The present study was made possible through a
research fellowship granted by the Treubel Found¬
ation (University of Basel, Switzerland) and research
funds of NIWA Oceanographic (Wellington). The
author wishes to express his sincere thanks to both
institutions for their generosity. Special thanks are
also due to Dr D.P. Gorcion (NIWA, Wellington), not
only for initiating anci supporting the present work.

but also for his friendly help in solving many other
problems. Many other people also contributed their
efforts to realise this work anci I wish to thank them
very much: Treffery Barnett (Aucklanci), all NIWA
staff in Wellington, Dr Jean Bouillon (Brussels), Dr
Dale Calcier (Toronto), Dr V. Schmid (Basel), Dr Wim
Vervoort (Leiden).

APPENDIX

Growth stages of Bougainvillia vervoorti

Meciusae were releaseci from polyps collected from nature. Several meciusae were examined but ciata are for one
representative medusa.

Day

Size

(mm)

Branching of
oral tentacles

Marginal
tentacles/ bulb

Remarks

0.8

tentacles short

tentacles grow

1.7

stomach base cruciform

1.8

2.2

manubrium turns green

2.6

perradial manubrium extensions grow

6-7

3.2

7-8

3.4

3.6

3-4

8-9

3.6

female gonad tissue well visible

Results of crosses between different morphs of Sarsia eximia

Pairs of medusae of Sarsia eximia belonging to both observed extrema of morphologies (normal bell-shape to cylindrical
form) were kept in small petri dishes in millipore-filtered sea water. The animals were kept for 16 hours in the dark
and then transferred to bright light which induced spawning.

The results are given below. The normal form is always left, the cylindrical form right. Not all sizes could be
measured because some bells shrunk overnight. In the columns labelled "manubrium lr , the length of the manubrium in
proportion to the bell cavity height is given. Only cross no. 4 did not yield larvae. The male used for this cross was
somewhat small and may have been immature. Otherwise, more than half of the spawneci eggs developed into embryos.

Legenci: sex = sex useci; size = umbrella height; manubrium = fraction of manubrium length to subumbrella height;
result = observed result, embryos were followed up to gastrula stage.

Rouncied bell Cylincirical bell Results

cross

sex

size

manubrium

sex

size

manubrium

male

1/2

female

i/i

10 embryos

female

3 mm

2/3

male

3.6 mm

i/i

30 embryos

female

1/1

male

i/i

11 embryos

male

2.2 mm

2/3

female

3.6 mm

2/3

few eggs, no development

male

3 mm

1/1

male

3.6 mm

1/1

17 embryos

144

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INDEX

The index covers the introduction, systematics and remarks.

Italic numbers indicate a text-figure; major references are indicated in bold type.

Acliaradria larynx 109
Amalda australis 46
Ampliinema 14, 62, 63
dinema 5, 62, 63 , 63, 64
rugosum 63, 64 , 65
Annatiara 62, 66, 66
ajfinis 66,66
Antarctica 42,43
Anthoathecata 6, 13, 84
Anthomedusa 6,7
Anthomedusae 90,139
Aphrodite 69
Argobiiccinum tumida 46
ArfennYi 9, 25, 36, 80, 87, 90, 92, 93,
96,110, 111, 118, 123, 135
Ascidioclava parasitica 5, 23
Aselomaris 40, 41, 42
arenosa 42

Aselomaris sagamiense 42
Asi/;/con/«c 92
Athecata 6, 7, 78
Athecatae 6
Auckland Harbour 143
Auckland Islands 31
Atractylis antarctica 41
Austrofiisus glans 46

Banks Peninsula 72
Bamettia 5, 62, 75

caprai 5, 75, 75, 76

Bimeria australis 13
Boeromedusa 115, 115
auricogonia 115
Boeromedusidae 115
Boltenia pachydermatina 23
Bougainvillia 27, 28, 32, 33, 36, 42
aurantiaca 27,28, 28
britannica 33

dimorpha 5, 27, 32, 37 , 38, 39
firim 27, 28 , 29, 36
inaequalis 27, 29 , 29
macloviana 27, 30 , 30
muscoides 28
tfwsci/s 27, 31 , 32, 33, 38
platygaster 27, 33 , 33
pyramidata 38,39
ramosa 31, 33
vervoorti 5, 27, 34 , 34, 35, 36,

38, 144

Bougainvilliidae 27, 40, 62
Bouillonia 107
Brancliiocerianthus 101
Bryozoa 95

Bytliotiara 22 , 22
huntsmani 23
murrayi 22,22
parasitica 5, 22, 23 , 24, 25
stilbosa 23
sp. 22, 25

Calycopsidae 22
Calycopsis 22, 25, 26
bigelowi 5, 25, 25, 26, 27
borchgrevinki 27
Calycopsis typa 25
Campbell Island 31
Candelabridae 118
Candelabrum 118
cocksii 118
sp. 118,218
Cape Foulwind 37,116
Cape Palliser 27, 72, 78
Capitata 90,139
Carpophyllum masclialocarpum 80
Chatham Islands 44
Cliitina 5,139,141
ericopsis 6,139,141
Cirrlii tiara 76

Cladocoryne 96

floccosa 5, 96, 97 , 97
Claoocorynidae 96
Cladonema 131 , 133

californicum 131,132,133
mayeri 131,133
novae-zelandiae 5, 131, 132
pacificum 131,133
perkinsi 131

radiatum 5,125, 131 , 232, 133
uchidai 133

Cladonematidae 131,133, 139
Clavidae 13, 62
Climacocodon 116
C//o cuspidata 72
Cnidonema capensis 133
liodgsoni 133
vallentini 133, 134
Cook Strait 25, 29, 33, 69, 70, 80,
128

Cominella maculosa 53
Congiopodus leucopaecilus 48
Coral lina 36

Cordylopliora 14
caspia 14,15
lacustris 15

lacustris var. otagoensis 15
rubra 5,18, 19

157

Corydendrium 15
zelandicum 5, 18,19
Corymorplia 102 , 102,105
forbesii 102 , 202, 103
furcata 102, 103 , 204
intermedia 5 , 102, 104 , 105
nutans 102,105
CORYMORPHIDAE 89, 101
Coryne 119 , 125, 12
cAimirt 125
muscoides 121,122
japonica 128
pusilla 5,119, 220

tenella 5, 125, 128

tricycla 5,119, 120 , 222, 122
vaginata 120,122
sp. 1 122 , 222,139

sp. 2 123 , 123

CORYNIDAE 5, 119, 139
Craspedacusta 6
Cricophorus nutrix 138
Ctenaria 92
ctenophora 92
Cubozoa 90
Cunina sp. 36
Cyanea macloviana 30
Cystophora 90,138
retroflexa 135
torulosa 135
Cytaeididae 61
Cytaeis 61 , 89
minima 50
octopunctata 59
tetrastyla 61 , 62, 62

Dianea conica 71
Dicoryne 27,39
annulata 40
conferta 39, 40
conybearei 5, 39 , 39, 40
Dipurena 6, 119, 123 , 125
ophiogaster 123 , 224, 125, 131
reesi 125
strangulata 123
sp. 89

Doubtful Sound 33
Dunedin 48
Dysmorphosa minuta 50

Ectopleura 107, 112

cracert 5,107, 208, 109, 110, 120
dumortieri 107
exxonia 107

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Ectopleura
jnponicn 10 7

larynx 107,109, 109, 110, 112
marina 107
mayeri 112
minerva 112

niulticirrata 5,107, 110, 111
pacifica 112
radiata 107
ralphi 107, 109
venusta 107
wrighti 109
sp. 107,112, 112
Eleutheria vallentini 133
Eleutheridae 133
Endocrypta huntsniani 23
parasitica 23
Eucodoniidae 5, 88, 90
Eucodonium 89, 89,101
arctica 89
broumei 5, 89, 89, 90
Eudendriidae 6,78
Eudendrium 78
insigne 80

maorianus 5, 78, 83, 83, 84
novaezelandiae 79, 79, 80
ramosum 31,78
ritchiei 79, 80, 81, 84
terrnnovae 79, 82, 82
sp. 82
Eugotoea 101
Euphysa 101, 106
aurata 106
japonica 128

problematica 5, 101,106,106
Eiipliysomma 106
Euphysora 101,102
flammed 106
fnrcata 103
japonica 106
Evans Bay 39,123,131

Fabienna 5, 84, 87
oligonema 5
sphaerica 5, 84, 86, 86, 87
Falkland Islands 59
Fenestrulina thyreophora 56
Fiordland 33, 56, 72

Fiordlandia 5, 43, 56

protecta 5, 56, 57
FukauraJiydra 101

Garveia 41
Geryonia octona 68
Gotoea 101

27, 40, 41

antarcticum 40, 41, 41, 42
Greta Point 9, 85
Gynmogonos 101

Haliterella sp. 76

Haliterella sp. 76
HalitJiolus 62, 67, 67, 68
cirratiis 67
intermedins 67,68
pwper 5, 67, 67, 68
Hali tiara 76
formosa 76
inflexa 77, 77, 78
Halocordyle 142
disticha 142
Halocoryne 6, 92, 93
epizoica 92
Hansiella 43
hermit crab 46, 48, 51, 53
Heteractis 106
Heterocordyle conybearei 39
Hippocrene 30
macloviana 30
platygaster 33
Hybocodon 107, 112
cryptus 113
forbesii 103
proh/er 82, 113, 113, 114

unicus 113, 114

Hydractinia 23, 43, 44, 45,46, 54,

echinata 46
lactea 43

novaezelandiae 5, 44, 45, 46
parvispma 44, 44, 45, 46
rubricata 5, 44, 46, 47, 48
HYDRAcriNirDAE 43,48
Hydroidomedusae 13
Hypolytus 106

Indonesia 27

Kaikoma 9, 37, 48, 84
Kantiella 87

enigma tica 85, 87

Kermadec Islands 7, 23,104

Kermadec Trench 62

Koellikeria 27, 42
fasciculata 42
maasi 42,42
Koellikerina 19; 27

Laingiomedusae 13,87
Leigh 9,33

Leigh Marine Reserve 28, 50, 59,
60, 69, 87, 88, 90,103,105,125
Leuckartiara 62, 67, 68, 69, 70
octona 67, 68, 69, 76
Limnomedusae 6,84
Lizzia formosissima 58

Lyttelton Harbour 30

Macrocystis pyrifera 19, 54, 97,128
Margelopsidae 116
Margelopsis 116
Medusa p or pit a 98

Medusa

velella 99
Meiorhopalon 105
Merga 62, 74, 75
treubeli 74, 75
violacea 74
Merona 13,21
Myrionema 78
Moerisiidae 90

Neoturris 62,69
papua 69, 70, 70
pileata 70
vesicaria 69
North Cape 30, 82

Oceania 14, 15
armata 15, 16
dinema 63
Octotiara 62
Oonautes 92
hanseni 92

Pagurus rubricatus 46, 48
Pandea 62, 70

conica 70, 71, 71, 72
Pandeidae 40, 62, 76

Pandeopsis 62, 72, 74

ikarii 72, 73, 73
scutigera 72
Parawrightia 40
Pelagohydra 116,117
mirabilis 6,116,117
Penion sp. 39
Pennaria 122, 142
australis 142,143
disfic/w 120, 142, 143
Pennariidae 142
Perigonimus antarcticum 41
maclovianus 30
serpens 63
yoldia-arcticae 67
Perigonymus muscus 31
Perna canalictdaris 110
Phialella 49
Platyhelminthes 6
Plotocnide 101, 106
borealis 89
Pochella 85

oligonema 5, 84, 85, 86
polynema 85

Podocoryna 43, 48, 50, 54, 89
australis 5, 48, 49, 50, 51, 52, 53,
54

bella 6, 48, 49
Crtmert 48, 53, 54
minima 48, 50, 50
minuta 48,50,52
selena 54
tenuis 51
Podocoryne 48, 51

158

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(cc )®®(S

Podocoryne ininuta 50
simplex 50
sp. 1 51

sp. 2 54
POLYORCHIDAE 90
Polyorchis penicillatus 59
Poor Knight Islands 142
Porpita 98,99
paciifica 98
porpita 98,98,99
umbella 98
PORPITIDAE 98

Portobello 9, 18, 25, 33, 46, 55 ,117
Proboscidactyla 84, 87, 88
abyssicola 87
flavicirrata 87
ornata 88

ornata van stolonifera 88
sp. 87, 88

Probosqdactylidae 6, 84
Protiaridae 76
Protiara formosa 76
Pteroclava 96
Pteropoda 62
PriLOCODIIDAE 13
Pyura pachydermatina 23, 54
rugata 23
sp. 41

Ralpharia 101, 107
Rathkea 58
formosissima 58, 59
lizzioides 59
octopunctata 58, 59, 60
Rathkeidae 58
Rhizodendrium 19,20
Rhizogeton 14,19, 21

conicum 5, 13,19, 20, 20, 21
ezoense 21
fusi forme 19

nudum 21
sterreri 19, 21
sp. 20,21,21
Rhizorhagiinae 40
Rhizorhagium 40,41
antarcticum 41
Rhizorhagium roseum 40

Rhynchozoon larreyi 93, 94, 95, 96

SrtrsiV? 6, 119, 125,127, 128

eximia 5, 125, 226, 127, 128, 130,
131,144

gracilis 125,127,128
japonica 5, 125,128, 129, 130,

131

ophiogaster 123
radiata 128
tubulosa 125
sp. 125

Siphonohydra 101
Siphonophora 13
Slabberia ophiogaster 123
Snares Islands 43
Solanderia 5, 139,141
ericopsis 5, 139, 140, 141
gracilis 139, 141
misakinensis 139, 141
secunda 141
SOLANDERJIDAE 139
Stauridiosarsia japonica 128
Staurocladia 131, 133, 138

lwdgsoni 133,134,135,138,139
vallentini 133, 234,135,138,139
wellingtoni 5, 133,135, 236, 237,
138,139

Steenstrupia 101,105
Stewart Island 7,43
Stomotoca atra 13
dinema 63
rugosa 64,65
Stylactaria 43, 44, 54, 55
betkensis 55
conchicola 55
inermis 54,55
otagoensis 5, 45,46, 54, 55
reticulata 55
SlYLASTERIDAE 6, 13
Symplectoscyphus johnstoni 56
Sy/iconi/Hp eximia 125
synonyms 15, 92,101,105,125

tentaculozooids 56
Terminology 7

Tiaranna affinis 66
ikarii 72
Tiari codon 90,122

coeruleus 90, 91, 92
sp. 90, 92, 92
Trachymedusae 90
Trichydra 85,87
pudica 85
Tubiclava 21
fructicosa 18, 19
rubra 5,18,19,128
Tubularia 107
attenoides 110
caspia 15
crocefl 107
larynx 109

mesembryanthemum 107
warreni 109
Tubulariidae 89,107,114
Turris papua 70
vesicaria 69
Turritopsis 12, 16, 49

nutricula 5, 16, 27, 18, 19, 128
pacifica 18, 19

Vannucccia 101, 102
forbesii 103
Velella 98, 99, 100
spirans 99
velella 99, 200, 101

Waitemata Harbour 3
Wellington Harbour 18, 25, 33, 36,
37, 40, 42, 64,4 67, 87, 92, 110, 111,
128

Whangateau Harbour 9, 76, 90,103,
106, 125

Zanclea 5, 93, 96
costata 93, 95, 96
polymorpha 5, 93, 94, 95, 96
Zancleais 93
Zancleidae 92
Z anclella 5, 92, 93, 95

bryozoophila 93
Zyzzyzus 107

159

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